Loudon, Andrew H; Woodhams, Douglas C; Parfrey, Laura Wegener; Archer, Holly; Knight, Rob; McKenzie, Valerie; Harris, Reid N
.... We describe the diversity of bacteria on red-backed salamanders (Plethodon cinereus) in the wild and the stability of these communities through time in captivity using culture-independent Illumina 16S rRNA gene sequencing...
Breanna L. Riedel; Kevin R. Russell; W. Mark Ford; Katherine P. O' Neill; Harry W. Godwin
Woodland salamander responses to either traditional grazing or silvopasture systems are virtually unknown. An information-theoretic modelling approach was used to evaluate responses of red-backed salamanders (Plethodon cinereus) to silvopasture and meadow conversions in southern West Virginia. Searches of area-constrained plots and artificial...
Eric B. Sucre; Jessica A. Homyack; Thomas R. Fox; Carola A. Haas
The use of amphibians as biological indicators of ecosystem health has received considerable attention because of the increasing importance placed upon maintaining biodiversity in forested ecosystems. In this study, we imposed three different eastern red-backed salamander (Plethodon cinereus) treatments: 1) low (n = 4; added 0 salamanders to each...
Siddig, Ahmed A; Ellison, Aaron M; Jackson, Scott
.... Such calibrations, however, are rare. The red back salamander, Plethodon cinereus, is an ecologically useful indicator species of forest dynamics, and accurate calibration of indices of salamander abundance could increase the reliability...
Sterrett, Sean; Brand, Adrianne; Fields, William R.; Katz, Rachel A.; Grant, Evan H. Campbell
Lungless salamanders (family Plethodontidae) are relatively sedentary and are presumed to have limited dispersal ability (Marsh et al. 2004. Ecology 85:3396–3405). Site fidelity in Plethodontidae is high, and individuals displaced 90 m return to home territories (Kleeberger and Werner 1982. Copeia 1982:409–415). Individuals defend territories (Jaeger et al. 1982. Anim. Behav. 30:490–496) and female home ranges have been estimated to be 24.34 m2 (Kleeberger and Werner 1982, op. cit.). Females may seek out suitable subsurface habitat to oviposit eggs, yet little is known about their maximum movement distances (Petranka 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington. 587 pp.).On 18 September 2014, a female P. cinereus (lead back morphotype; SVL = 44.68 mm; 0.89 g) was found under a coverboard during a standard sampling event and uniquely marked using visual implant elastomer at the S.O. Conte Anadromous Fish Research Center, Massachusetts, USA (42.59280°N, 72.58070°W, datum WGS84; elev. 74 m). This individual was subsequently recaptured at ~1500 h on 8 October 2014 under a coverboard within 3 m of the original capture location and then again ~1430 h on 16 October 2014 under a log, within the same forest patch, though in a 50 x 150 m area adjacent to the original study area. Because we found the marked salamander while collecting multiple individuals for a laboratory study, the exact recapture location of the marked individual is not known. However, the distance between the 8 October capture location and the nearest edge of the 16 October search area (i.e. 50 x 150 m) was 143 m, indicating a minimum movement distance. As far as we are aware, this is the longest recorded movement for P. cinereus by more than 53 m (Kleeberger and Werner 1982, op. cit.). This finding followed a rain event of 1.63 cm within 24 h and the second largest sustained rain event during October. The movement we observed may have been due to
Dawley, E M; Fingerlin, A; Hwang, D; John, S S; Stankiewicz, C A
The chemosensory epithelium of vertebrates retains the ability to produce new receptor neurons throughout life, presumably as a mechanism to replace aging or damaged receptors. We examined cell division in the main olfactory and vomeronasal epithelia of red-backed salamanders (Plethodon cinereus) because previous studies had shown that the volume of sensory epithelia changes seasonally. Cell division was compared throughout the year by injecting salamanders once with 5-bromo-2'-deoxyuridine (BrdU), which is incorporated into the DNA of cells during DNA synthesis, and sacrificing them one hour after injection. We used immunocytochemistry to locate cells that had arisen from cell division since BrdU injection and compared the number of labeled cells per area among animals. Animals collected in May had significantly more labeled nuclei than animals collected in any other month. However, proliferation rates among the other months were not significantly different and were quite low. Labeled nuclei also were found around the cerebral ventricles of salamanders collected in May, but rarely in any other month, although other tissues in the head often were heavily labeled. Cell proliferation appears to be up-regulated in the chemosensory epithelia and in the telencephalon during May, and we hypothesize that new receptors, and perhaps their interneurons in the telencephalon, are being generated in anticipation of seasonal events that are mediated by chemoreception. Copyright 2000 S. Karger AG, Basel
Breanna L. Riedel; Kevin R. Russell; W. Mark. Ford
Nonforested habitats such as open fields and pastures have been considered unsuitable for desiccation-prone woodland salamanders such as the Eastern Red-backed Salamander (Plethodon cinereus). Recent research has suggested that Plethodon cinereus may not only disperse across but also reside within open habitats including fields,...
Bazar, Matthew A; Quinn, Michael J; Mozzachio, Kristie; Bleiler, John A; Archer, Christine R; Phillips, Carlton T; Johnson, Mark S
Copper (Cu) has widespread military use in munitions and small arms, particularly as a protective jacket for lead projectiles. The distribution of Cu at many US military sites is substantial and sites of contamination include habitats in and around military storage facilities, manufacturing, load and packing plants, open burning/open detonation areas, and firing ranges. Some of these areas include habitat for amphibian species, which generally lack toxicity data for risk assessment purposes. In an effort to ascertain Cu concentrations in soil that are toxic to terrestrial amphibians, 100 red-backed salamanders (Plethodon cinereus) were randomly sorted by weight, assigned to either a control soil or one of four treatments amended with copper acetate in soil, and exposed for 28 days. Analytical mean soil concentrations were 18, 283, 803, 1333, and 2700 mg Cu/kg soil dry weight. Food consisted of uncontaminated flightless Drosophila melanogaster. Survival was reduced in salamanders exposed to 1333 and 2700 mg/kg by 55% and 100%, respectively. Mortality/morbidity occurred within the first 4 days of exposure. These data suggest that a Cu soil concentration of and exceeding 1333.3 +/- 120.2 mg/kg results in reduced survival, whereas hematology analyses suggest that a concentration of and exceeding 803.3 +/- 98.4 mg/kg might result in reduced total white blood cell count. No effects were observed at 283.3 +/- 36.7 mg/kg.
Loudon, Andrew H; Woodhams, Douglas C; Parfrey, Laura Wegener; Archer, Holly; Knight, Rob; McKenzie, Valerie; Harris, Reid N
Beneficial cutaneous bacteria on amphibians can protect against the lethal disease chytridiomycosis, which has devastated many amphibian species and is caused by the fungus Batrachochytrium dendrobatidis. We describe the diversity of bacteria on red-backed salamanders (Plethodon cinereus) in the wild and the stability of these communities through time in captivity using culture-independent Illumina 16S rRNA gene sequencing. After field sampling, salamanders were housed with soil from the field or sterile media. The captive conditions led to different trajectories of bacterial communities. Eight OTUs present on >90% of salamanders in the field, through time, and in both treatments were defined as the core community, suggesting that some bacteria are closely associated with the host and are independent of an environmental reservoir. One of these taxa, a Pseudomonas sp., was previously cultured from amphibians and found to be antifungal. As all host-associated bacteria were found in the soil reservoir, environmental microbes strongly influence host-microbial diversity and likely regulate the core community. Using PICRUSt, an exploratory bioinformatics tool to predict gene functions, we found that core skin bacteria provided similar gene functions to the entire community. We suggest that future experiments focus on testing whether core bacteria on salamander skin contribute to the observed resistance to chytridiomycosis in this species even under hygenic captive conditions. For disease-susceptible hosts, providing an environmental reservoir with defensive bacteria in captive-rearing programs may improve outcomes by increasing bacterial diversity on threatened amphibians or increasing the likelihood that defensive bacteria are available for colonization.
Female parent-offspring phenotypic data on color morph frequencies in the red-backed salamander, Plethodon cinereus, were obtained from two Virginia localities (164 broods from Greene County and 97 broods from Giles County). The color morph data indicate that the striped morph is genetically dominant in Giles County and recessive in Greene County. It is suggested that epistatic interaction of two or more loci is responsible for the difference between the localities.
Braekevelt, C R
The retinal pigment epithelium (RPE) of the red-backed salamander (Plethodon cinerus) consists of a single layer of large squamous shaped cells. The RPE cells are but minimally infolded basally (sclerally) but show many large apical (vitreal) processes interdigitating with the rod outer segments. These epithelial cells are joined laterally by prominent tight junctions located in the mid region of the cells. Internally smooth endoplasmic reticulum is very plentiful while rough endoplasmic reticulum is not. Polysomes, small dense mitochondria and small round to oval melanosomes are plentiful. Golgi zones and lysosome-like bodies are also present as are phagosomes of outer segment material and myeloid bodies. The RPE cell nucleus is large and vesicular. It is felt that the melanosomes undergo retinomotor movements but as only light-adapted specimens were examined it is not known how extensive are these movements. Bruch's membrane or complexus basalis shows the typical pentalaminate structure noted for most vertebrates. The choriocapillaris is a single layer of large anastomosing capillaries which are minimally fenestrated facing Bruch's membrane.
Braekevelt, C R
The retinal photoreceptors of the red-backed salamander (Plethodon cinerus) have been studied by light and electron microscopy. Rods and single cones are present in this duplex retina in a ratio of about 25:1. The photoreceptors in this amphibian species are much larger than is reported for most vertebrates. In the light-adapted state, rods reach deep into the retinal epithelial (RPE) layer. The rod outer segment is composed of discs of uniform diameter displaying several very deep incisors. The rod inner segment displays a distal elliposid of mitochondria and a short stout myoid region. Rod nuclei are electron dense and often protrude through the external limiting membrane. Rod synaptic spherules are large and display several invaginated synaptic sites as well as superficial synapses. It is felt that the rods do not undergo retinomotor movements. The cone photoreceptors are much smaller than the rods and display a tapering outer segment, an unusual modified ellipsoid and a large parabolid of glycogen in the inner segment. Cone nuclei are less electron dense than rods and are located at all levels within the outer nuclear layer. The synaptic pedicle of the cones is larger, more electron lucent and display more synaptic sites (both invaginated and superficial) than that of rods. It is felt that cone photomechanical responses are minimal.
Dawley, E M; Crowder, J
Sexually dimorphic behaviors often are associated with sexually dimorphic neural structures. Perinatal hormonal levels influence structural sexual dimorphism, and seasonal structural changes also can be the result of seasonal hormonal fluctuations. We compared the volume of vomeronasal organs of male and female red-backed salamanders (Plethodon cinereus) throughout the year. Odorants are delivered to vomeronasal receptors through nasolabial grooves when salamanders touch the bases of these grooves to objects (nose tapping). Males may locate and identify potential mates through nose tapping during the prolonged breeding season that lasts from October through May. We compared vomeronasal organ data through multiple regression by using total body size, sex, and season as variables that may influence vomeronasal organ volume. Gonads also were examined as an indicator of reproductive status. Total body size and sex significantly affect vomeronasal organ volume; as body size increases, so does vomeronasal organ volume, and males have significantly larger vomeronasal organs than females at all times of the year. During the summer, both males and females have larger vomeronasal organs than at any other time of the year. Summer also is a period of intense foraging and the initiation of a new cycle of gametogenesis. Previous studies of circulating hormone levels in amphibians have shown that the initial stages of gametogenesis correspond to a period of low estradiol and testosterone levels but high gonadotropin-releasing hormone levels. The functional significance of sexual and seasonal differences in the vomeronasal organs of P. cinereus may relate to the neurogenesis of specialized receptors for courtship and mating.
Bondi, Cheryl A; Beier, Colin M.; Ducey, Peter K; Lawrence, Gregory B.; Bailey, Scott W.
Hardwood forests of eastern North America have experienced decades of acidic deposition, leading to soil acidification where base cation supply was insufficient to neutralize acid inputs. Negative impacts of soil acidity on amphibians include disrupted embryonic development, lower growth rates, and habitat loss. However, some amphibians exhibit intraspecific variation in acid tolerance, suggesting the potential for local adaptation in areas where soils are naturally acidic. The eastern red-backed salamander (Plethodon cinereus) is a highly abundant top predator of the northern hardwood forest floor. Early research found that P. cinereus was sensitive to acidic soils, avoiding substrates with pH < 3.8 and experiencing decreased growth rates in acidic habitats. However, recent studies have documented P. cinereus populations in lower pH conditions than previously observed, suggesting some populations may persist in acidic conditions. Here, we evaluated relationships between organic horizon soil pH and P. cinereus abundance, adult health (body size and condition), and microhabitat selection, based on surveys of 34 hardwood forests in northeastern United States that encompass a regional soil pH gradient. We found no associations between soil pH and P. cinereus abundance or health, and observed that this salamander used substrates with pH similar to that available, suggesting that pH does not mediate their fine-scale distributions. The strongest negative predictor of P. cinereus abundance was the presence of dusky salamanders (Desmognathus spp.), which were most abundant in the western Adirondacks. Our results indicate that P. cinereus occupies a wider range of soil pH than has been previously thought, which has implications for their functional role in forest food webs and nutrient cycles in acid-impaired ecosystems. Tolerance of P. cinereus for more acidic habitats, including anthropogenically acidified forests, may be due to local adaptation in
Riedel, Breanna L.; Russell, Kevin R.; W. Mark Ford
Nonforested habitats such as open fields and pastures have been considered unsuitable for desiccation-prone woodland salamanders such as the Eastern Red-backed Salamander (Plethodon cinereus). Recent research has suggested that Plethodon cinereus may not only disperse across but also reside within open habitats including fields, meadows, and pastures. However, presence and high densities of P. cinereus within agriculturally disturbed habitats may be misleading if these populations exhibit aty...
Telfer, A C; Laberge, F
Terrestrial salamanders are able to detect prey items using chemical cues, but the nature of the cues involved is uncertain. This study aimed to tease apart the roles of the soluble and volatile components of prey cues detected by Eastern red-backed salamanders (Plethodon cinereus), assuming the likelihood that these different components are respectively detected by the vomeronasal (accessory) and main olfactory organs. Wild-caught salamanders were exposed to control or soluble and volatile cricket cues in two different behavioural assays conducted in the laboratory. The first series of assays focused on localized presentation of soluble cues on the substrate, and the second on point sources of volatile cues delivered through plastic tubes. Room temperature was varied across experiments. Salamanders increased chemoinvestigation of the substrate via nosetapping when soluble prey cues were distributed non-uniformly on the substrate. In the warmer of two temperatures tested, salamanders additionally showed a spatial preference for location of soluble cue deposition. Attraction to a point source of volatile cues was not evident when examining the responses of salamanders grouped together; however, investigation of the volatile point source was significantly correlated with side preference only when both soluble cues and a volatile point source were present. The latter suggests that a subset of salamanders were attracted to the point source of volatile cues in the presence of soluble cues on the substrate. This study indicates that soluble prey cues alone are sufficient to trigger salamander foraging behaviour, and that temperature influences this foraging response. It supports the notion that the vomeronasal system plays an important role in prey detection, but suggests that volatile cues are also investigated by some salamanders when soluble prey cues have been detected.
Johnson, Mark S. [U.S. Army Center for Health Promotion and Preventive Medicine, Health Effects Research Program, 5158 Blackhawk Road, ATTN: MCHB-TS-THE, Aberdeen Proving Ground, MD 21010-5403 (United States)]. E-mail: firstname.lastname@example.org; Suski, Jamie [U.S. Army Center for Health Promotion and Preventive Medicine, Health Effects Research Program, 5158 Blackhawk Road, ATTN: MCHB-TS-THE, Aberdeen Proving Ground, MD 21010-5403 (United States); Bazar, Matthew A. [U.S. Army Center for Health Promotion and Preventive Medicine, Health Effects Research Program, 5158 Blackhawk Road, ATTN: MCHB-TS-THE, Aberdeen Proving Ground, MD 21010-5403 (United States)
Dinitrotoluenes are used as propellants and in explosives by the military and as such have been found at relatively high concentrations in the soil. To determine whether concentrations of 2,4-dinitrotoluene (2,4-DNT) in soil are toxic to amphibians, 100 red-backed salamanders (Plethodon cinereus) were exposed to either 1500, 800, 200, 75 or 0 mg 2,4-DNT/kg soil for 28 days and evaluated for indicators of toxicity. Concentrations of 2,4-DNT were less than targets and varied with time. Most salamanders exposed to concentrations exceeding 1050 mg/kg died or were moribund within the first week. Salamanders exposed to soil concentrations exceeding 345 mg/kg lost >6% of their body mass though no mortality occurred. Overt effects included a reduction in feed consumption and an increase in bucco-pharyngeal oscillations in salamanders. These results suggest that only high soil concentrations of 2,4-DNT have the potential to cause overtly toxic effects in terrestrial salamanders. - Exposures of 2,4-dinitrotoluene in soil exceeding 345 mg/kg causes toxicity to P. cinereus.
Johnson, Mark S; Suski, Jamie; Bazar, Matthew A
Dinitrotoluenes are used as propellants and in explosives by the military and as such have been found at relatively high concentrations in the soil. To determine whether concentrations of 2,4-dinitrotoluene (2,4-DNT) in soil are toxic to amphibians, 100 red-backed salamanders (Plethodon cinereus) were exposed to either 1500, 800, 200, 75 or 0mg 2,4-DNT/kg soil for 28 days and evaluated for indicators of toxicity. Concentrations of 2,4-DNT were less than targets and varied with time. Most salamanders exposed to concentrations exceeding 1050 mg/kg died or were moribund within the first week. Salamanders exposed to soil concentrations exceeding 345 mg/kg lost >6% of their body mass though no mortality occurred. Overt effects included a reduction in feed consumption and an increase in bucco-pharyngeal oscillations in salamanders. These results suggest that only high soil concentrations of 2,4-DNT have the potential to cause overtly toxic effects in terrestrial salamanders.
McKenny, H.C.; Keeton, W.S.; Donovan, T.M.
Managing for stand structural complexity in northern hardwood forests has been proposed as a method for promoting microhabitat characteristics important to eastern red-backed salamanders (Plethodon cinereus). We evaluated the effects of alternate, structure-based silvicultural systems on red-backed salamander populations at two research sites in northwestern Vermont. Treatments included two uneven-aged approaches (single-tree selection and group-selection) and one unconventional approach, termed "structural complexity enhancement" (SCE), that promotes development of late-successional structure, including elevated levels of coarse woody debris (CWD). Treatments were applied to 2 ha units and were replicated two to four times depending on treatment. We surveyed red-backed salamanders with a natural cover search method of transects nested within vegetation plots 1 year after logging. Abundance estimates corrected for detection probability were calculated from survey data with a binomial mixture model. Abundance estimates differed between study areas and were influenced by forest structural characteristics. Model selection was conducted using Akaike Information Criteria, corrected for over-dispersed data and small sample size (QAICc). We found no difference in abundance as a response to treatment as a whole, suggesting that all of the uneven-aged silvicultural systems evaluated can maintain salamander populations after harvest. However, abundance was tied to specific structural habitat attributes associated with study plots within treatments. The most parsimonious model of habitat covariates included site, relative density of overstory trees, and density of more-decayed and less-decayed downed CWD. Abundance responded positively to the density of downed, well-decayed CWD and negatively to the density of poorly decayed CWD and to overstory relative density. CWD volume was not a strong predictor of salamander abundance. We conclude that structural complexity enhancement
Bazar, Mathew A; Quinn, Michael J; Mozzachio, Kristie; Johnson, Mark S
Since World War I, trinitrotoluene (TNT) has been the most commonly used explosive. Environmental contamination associated with synthesis, manufacture of weapons, and use during training exercises has been extensive, with soil concentrations reaching 145,000 mg/kg. Some of these areas include habitats for amphibian species. Earlier studies have shown that salamanders dermally absorb TNT from soil. To ascertain what soil concentrations of TNT are toxic to amphibians, red-backed salamanders (Plethodon cinereus) were exposed to one of five concentrations of TNT in soil for 28 d and evaluated for indicators of toxicity. A total of 100 salamanders were randomly sorted by weight and assigned to target TNT concentrations in soil of either 3,000, 1,500, 750, 325, or 0 mg/kg dry weight. Food consisted of uncontaminated flightless Drosophila melanogaster. Survival was reduced in salamanders exposed to 1,500 and 3,000 mg/kg by 10 and 55%, respectively. Most mortality/morbidity occurred within the first week of exposure. Salamanders had a reduction in hemoglobin at 750 mg/kg or greater and a reduction in red blood cell concentration at 1,500 mg/kg or greater. Food consumption was affected in salamanders at 750 mg/kg or greater; a reduction in body mass and liver glycogen content also occurred at and above this concentration. Splenic congestion also was observed in salamanders from these groups. These data suggest that soil TNT concentrations of 373 +/- 41.0 mg/kg or greater result in reduced body mass, reduced feed intake, and hematological effects.
Dinsmore, C E; Hanken, J
Species of the salamander genus Plethodon have a characteristically uniform morphology. Morphological conservation at the level of interspecific comparisons, however, is not always reflected within species. Perhaps the most extreme example of intraspecific variation is the recent description of extensive variability in limb-skeletal patterning both within and between populations of the widespread species P. cinereus. We utilized limb regeneration following experimental amputation as a tool to examine whether naturally occurring variant skeletal patterns result from limb loss and regeneration in nature, and to assay the intrinsic (i.e., genetic) component of between-individual variation in mesopodial patterning. We observed the following. First, regenerate patterns are strikingly different from native patterns: interelement fusions in regenerates are typically between proximodistally adjacent cartilages, whereas interelement fusions in native variant limbs occur exclusively between laterally adjacent cartilages. Fusions also are over ten times more frequent in regenerates than in native limbs. Second, there is no strong correlation between native limb pattern (typical vs. variant) and the regenerate pattern. We conclude that variability in field-collected P. cinereus reflects extensive intrapopulation variation in limb-skeletal patterning during original limb development, rather than regeneration in nature, and that limb regeneration analysis provides no evidence of a strong genetic component to between-individual variation. Finally, unusual mesopodial patterns produced during limb regeneration may be related to the mechanical factors impinging on the regenerating limb in this terrestrial species.
Davis, Andrew K; DeVore, Jayna L; Milanovich, Joseph R; Cecala, Kristen; Maerz, John C; Yabsley, Michael J
During a recent study of red-backed salamanders (Plethodon cinereus), we discovered an intraerythrocytic organism typified by violet-staining, intracellular inclusions, consistent with descriptions of Cytamoeba or Aegyptianella (bacteria). Here we characterize its taxonomic status using molecular techniques and ask basic questions about its nature. Blood smears from 102 salamanders were examined from Pennsylvania, New York, and Virginia to determine prevalence, and whole blood from several infected animals was tested using a PCR which targets the 16S rRNA gene of bacteria. Phylogenetic analysis of partial 16S rRNA gene sequence (1201 bp) indicated this organism was in the order Rickettsiales and is likely a member of the family Anaplasmatacea. The organism differed from currently described taxa and was clearly differentiated from Aegyptianella pullorum of birds and "Candidatus Hemobacterium ranarum" (formally A. ranarum) of frogs. Of all salamanders, 17 (16.7%) were infected and these were significantly larger (snout-vent length) and had higher body condition scores than uninfected ones, and males were more likely to be infected than females. Erythrocytes affected by the pathogen were 5% larger than unaffected ones, but otherwise similar in morphology. Infected animals tended to have a greater number of circulating white blood cells, based on estimates from smears, indicating a nonspecific response to the pathogen by the innate immune system. Given its phylogenetic position, this pathogen is likely transmitted by an arthropod vector, and the male-biased prevalence strongly implicates trombiculid mites, which also live in leaf litter and affect male salamanders more so than females.
Ahmed A. Siddig
Full Text Available Herpetologists and conservation biologists frequently use convenient and cost-effective, but less accurate, abundance indices (e.g., number of individuals collected under artificial cover boards or during natural objects surveys in lieu of more accurate, but costly and destructive, population size estimators to detect and monitor size, state, and trends of amphibian populations. Although there are advantages and disadvantages to each approach, reliable use of abundance indices requires that they be calibrated with accurate population estimators. Such calibrations, however, are rare. The red back salamander, Plethodon cinereus, is an ecologically useful indicator species of forest dynamics, and accurate calibration of indices of salamander abundance could increase the reliability of abundance indices used in monitoring programs. We calibrated abundance indices derived from surveys of P. cinereus under artificial cover boards or natural objects with a more accurate estimator of their population size in a New England forest. Average densities/m2 and capture probabilities of P. cinereus under natural objects or cover boards in independent, replicate sites at the Harvard Forest (Petersham, Massachusetts, USA were similar in stands dominated by Tsuga canadensis (eastern hemlock and deciduous hardwood species (predominantly Quercus rubra [red oak] and Acer rubrum [red maple]. The abundance index based on salamanders surveyed under natural objects was significantly associated with density estimates of P. cinereus derived from depletion (removal surveys, but underestimated true density by 50%. In contrast, the abundance index based on cover-board surveys overestimated true density by a factor of 8 and the association between the cover-board index and the density estimates was not statistically significant. We conclude that when calibrated and used appropriately, some abundance indices may provide cost-effective and reliable measures of P. cinereus
Bazar, Matthew A; Quinn, Michael J; Mozzachio, Kristie; Bleiler, John A; Archer, Christine R; Phillips, Carlton T; Johnson, Mark S
The use of lead in military and civilian small arms projectiles is widely acknowledged to have resulted in high soil lead concentrations at many small arms ranges. These ranges are often adjacent to wildlife habitat or have become habitat when no longer used. To assess the potential toxicity of lead to terrestrial amphibians in contaminated areas, we exposed 100 red-backed salamanders (Plethodon cinereus) to either a control soil or one of four soil treatments amended with lead acetate for 28 days. Analytical mean soil concentrations were 14 (control), 553, 1700, 4700, and 9167 mg Pb/kg soil dry weight. An additional 60 salamanders were also exposed for 28 days to one of six field-collected soil samples from a small arms range and a skeet range. The field soil concentrations ranged from 11 (background) to 16,967 mg Pb/kg soil dry weight. Food consisted of uncontaminated flightless Drosophila melanogaster. Salamander survival was reduced in amended soil treatments of 4700 and 9167 mg/kg by 15% and 80%, respectively. Inappetence was observed at 4700 and 9167 mg/kg and growth decreased in the 9167 mg/kg treatment. Total white blood cells decreased 32% at 4700 mg/kg compared to controls and were 22% lower in the 9167 mg/kg treatment. In contrast, survival was 100% for all field-collected soils with no hematological effects. At 16,967 mg/kg there was evidence of soil avoidance and decreased growth. These data suggest marked differences in toxicity and bioavailability of the lead-amended soil in contrast to the field-collected soil containing lead.
Siddig, Ahmed A; Ellison, Aaron M; Jackson, Scott
Herpetologists and conservation biologists frequently use convenient and cost-effective, but less accurate, abundance indices (e.g., number of individuals collected under artificial cover boards or during natural objects surveys) in lieu of more accurate, but costly and destructive, population size estimators to detect and monitor size, state, and trends of amphibian populations. Although there are advantages and disadvantages to each approach, reliable use of abundance indices requires that they be calibrated with accurate population estimators. Such calibrations, however, are rare. The red back salamander, Plethodon cinereus, is an ecologically useful indicator species of forest dynamics, and accurate calibration of indices of salamander abundance could increase the reliability of abundance indices used in monitoring programs. We calibrated abundance indices derived from surveys of P. cinereus under artificial cover boards or natural objects with a more accurate estimator of their population size in a New England forest. Average densities/m(2) and capture probabilities of P. cinereus under natural objects or cover boards in independent, replicate sites at the Harvard Forest (Petersham, Massachusetts, USA) were similar in stands dominated by Tsuga canadensis (eastern hemlock) and deciduous hardwood species (predominantly Quercus rubra [red oak] and Acer rubrum [red maple]). The abundance index based on salamanders surveyed under natural objects was significantly associated with density estimates of P. cinereus derived from depletion (removal) surveys, but underestimated true density by 50%. In contrast, the abundance index based on cover-board surveys overestimated true density by a factor of 8 and the association between the cover-board index and the density estimates was not statistically significant. We conclude that when calibrated and used appropriately, some abundance indices may provide cost-effective and reliable measures of P. cinereus abundance that
Full Text Available Hemlock forests of the northeastern United States are declining due to the invasive hemlock woolly adelgid (HWA (Adelges tsugae. Hardwood species replace these forests, which affects soil properties that may influence other communities, such as red-backed salamanders (red-backs (Plethodon cinereus. This study examined the effects of HWA invasion on soil properties and how this affects red-backs at the Hemlock Removal Experiment at Harvard Forest, which consists of eight 0.8 ha plots treated with girdling to simulate HWA invasion, logging to simulate common management practices, or hemlock- or hardwood-dominated controls. Coverboard surveys were used to determine the relative abundance of red-backs between plots during June and July 2014 and soil cores were collected from which the bulk density, moisture, pH, temperature, leaf litter, and carbon-nitrogen ratio were measured. Ordination provided a soil quality index based on temperature, pH, and carbon-to-nitrogen ratio, which was significantly different between plot treatments (p < 0.05 and showed a significant negative correlation with the red-back relative abundance (p < 0.05. The findings support the hypothesis that red-backs are affected by soil quality, which is affected by plot treatment and thus HWA invasion. Further studies should explore how salamanders react in the long term towards changing environments and consider the use of red-backs as indicator species.
Simons, R R; Felgenhauer, B E; Thompson, T
Plethodon cinereus, the red-backed salamander, is a small territorial vertebrate that defends refugia located on the forest floor. As a component of territorial defense, these animals use scent marks to advertise their refugia. Behavioral evidence indicates that scent marks are produced by the postcloacal glands located on the ventral surface of the tail just posterior to the cloaca. We placed animals on unmarked substrates and recorded changes in serous acini from the postcloacal and shoulder region over a 48-h period. Within the first hour there was an increase in the number of acini filled with secretory product in the postcloacal region. At 12 h the number of full acini decreased and the number of empty acini increased. By 24 h the number of empty acini had decreased and the number of renewing acini containing secretory cells producing product had increased. By 48 h the ratio of full to empty to renewing acini was similar to those observed at the start of the study. In the shoulder region, no significant changes in the ratio of full to empty to renewing acini were observed. Observations of the serous acini within the postcloacal region and the shoulder region indicate that the mode of secretory production is holocrine. These findings are additional evidence that the postcloacal glands are the site of scent mark production. Copyright 1999 Wiley-Liss, Inc.
Sever, D M
In northern Indiana, the mating season of Plethodon cinereus occurs after hibernation from March until June, when oviposition begins. During the mating season, a female stores sperm in its spermatheca, a compound tubular gland in the roof of the cloaca. The apical cytoplasm of the spermathecal epithelium is filled with large secretory vacuoles whose product is released while sperm are stored. Females induced to oviposit in June and July by injections of human chorionic gonadotropin (hCG) still retain much sperm 1 month after oviposition, but secretory vacuoles are absent in all specimens sacrificed in July and August. Instead, some sperm are embedded in the spermathecal epithelium with resultant spermiophagy involving lysosomes. A female sacrificed in September 2 months after oviposition possesses scant sperm, but spermiophagy alone does not seem extensive enough to account for the decrease in sperm numbers. Females sacrificed in October prior to hibernation lack sperm in their spermathecae; some secretory vacuoles are present, but they are not as numerous or as enlarged as in specimens collected in March and May. Inter- and intrafamilial differences in the cytology of sperm storage may not be phyletically informative at the family level but related to species-specific reproductive adaptations.
Breanna L. Riedel
Full Text Available Nonforested habitats such as open fields and pastures have been considered unsuitable for desiccation-prone woodland salamanders such as the Eastern Red-backed Salamander (Plethodon cinereus. Recent research has suggested that Plethodon cinereus may not only disperse across but also reside within open habitats including fields, meadows, and pastures. However, presence and high densities of P. cinereus within agriculturally disturbed habitats may be misleading if these populations exhibit atypical demographic characteristics or decreased physical condition relative to forest populations. We surveyed artificial cover boards from 2004-2005 to compare physical condition, sex ratios, and age-class structure of P. cinereus among woodland, woodland-meadow edge, silvopasture, and meadow sites in the central Appalachian Mountains of southern West Virginia. Physical condition of salamanders was not significantly different among the four habitats. Furthermore, adult sex ratios of P. cinereus typically were not significantly different from 1 : 1 and were similar between forested and non-forested sites, although populations within silvopastures were biased towards females. However, adult salamanders were significantly more abundant than juveniles in all habitat types, with differences most pronounced within meadow habitats. Our study indicates that relatively small, non-forested habitats such as silvopastures and meadows may not adversely affect the overall physical condition or sex ratios of Plethodon cinereus. However, the paucity of juveniles within disturbed meadows indicates that these agriculturally modified habitats may negatively impact reproduction or that immature salamanders are at a physiological or competitive disadvantage in comparison to adults, particularly when surface cover is limited.
Sever, David M; Siegel, Dustin S
Caudal courtship glands (CCGs) are sexually dimorphic glands described in the skin of the dorsal tail base of some male salamanders in the genera Desmognathus, Eurycea, and Plethodon in the family Plethodontidae. These glands are believed to deliver pheromones to females during courtship, when the female rests her chin on the dorsal tail base during the stereotypic tail straddling walk unique to plethodontids. Although CCGs have been studied histologically, no investigations of their ultrastructure have been made. This article presents the first study on the fine structure and seasonal variation of CCGs, using the plethodontid Plethodon cinereus. The CCGs vary seasonally in height and secretory activity. The mature secretory granules observed in males collected in October and April consist of oval, biphasic granules that are eosinophilic and give positive reactions to periodic acid-Schiff for neutral carbohydrates but do not stain for acidic mucosusbtances or proteins with alcian blue and bromphenol blue, respectively. Granular glands, some of which contain mucous demilunes, are twice as large as CCGs, are syncytial (unlike CCGs), and stain for proteins. Mucous glands are similar in size to CCGs, but are basophilic, show no seasonal variation in secretory activity, and stain positive for acidic mucosubstances. CCGs do not resemble cytologically the sexually dimorphic mental glands of some plethodontids, which contain round or oval granules filled with an electron-dense amorphous substance. The CCGs are similar histologically to sexually dimorphic skin glands described in some anurans, but more comparative work is needed. © 2014 Wiley Periodicals, Inc.
Dawley, Ellen M; Nelsen, Meaghan; Lopata, Adrianne; Schwartz, Jessica; Bierly, Alison
In addition to the continuous low levels of neurogenesis typical of adult vertebrates to replace damaged chemoreceptor cells, red-backed salamanders (Plethodon cinereus) experience an up-regulation of chemoreceptor epithelial cell proliferation on a seasonal basis. Significantly more cell division occurs in late spring than at any other time of the year, and we investigated the fate and life span of these newly generated cells. We used 5-bromo-2'-deoxyuridine (BrdU) immunocytochemical cell birth dating to examine cell proliferation and cell migration in the main olfactory and vomeronasal epithelia of red-backed salamanders collected in late spring who were allowed to survive for one hour or three, four, 25, 28, 42, 49 and 100 days post-injection. We examined new neuron growth in the vomeronasal and olfactory epithelia using antibodies against Growth Associated Protein-43 (GAP-43), a protein whose synthesis is up-regulated during axon growth. We also tracked apoptosis within both types of chemosensory epithelia using terminal deoxynucleotidyl transferase-mediated nick-end labeling (TUNEL). BrdU-immunoreactive cells were located extensively throughout the vomeronasal epithelia, particularly in the area posterior to the entrance of the nasolacrimal duct, not only after three days of survival, but also all of the longer experimental survival periods as well; BrdU-ir cells within the olfactory epithelia were rarely located after longer survival periods. Salamanders collected in late spring displayed extensive GAP-43 labeling in the vomeronasal epithelia posterior to the entrance of the nasolacrimal duct, indicating a large population of young vomeronasal receptor neurons. Finally, apoptotic cells were evident in this same post-nasolacrimal-duct-area of the vomeronasal organ and in the olfactory epithelium. We suggest that vomeronasal receptor neurons born in late spring function throughout the summer and may be associated with the animals' extensive territoriality
Andrew K. DAVIS, Joseph R. MILANOVICH
Full Text Available Throughout the animal kingdom there are species that have two or more phenotypic forms or ‘morphs’, and many of these are amphibians. In North America, the red-backed salamander Plethodon cinereus can have either a red dorsal stripe or no dorsal stripe (lead-phase form, and evidence to date indicates the lead-phase form incurs a greater number of attacks from predators. In a recent collection of 51 P. cinereus, blood smears of both color morphs (35 red-stripe, 16 lead-phase were examined to obtain numbers of circulating leukocytes (via light microscopy, which can be used to indirectly estimate levels of stress hormones in vertebrates via a ‘hematological stress index’, which is the ratio between the number of two leukocyte types (neutrophils and lymphocytes. Our results showed that lead-phase salamanders tended to have greater numbers of circulating neutrophils and lower numbers of circulating lymphocytes than red-stripe morphs, leading to higher average neutrophil-lymphocyte ratios in lead-phase individuals. Since the salamanders were held (refrigerated for 7 days before sampling, we cannot be certain if this effect is a stress reaction to the captivity or the normal level for this morph. However comparison with two sets of related salamanders that were captured and sampled immediately indicates the red-stripe salamanders were either not stressed from the captivity at all, or their white blood cell distributions had returned to normal after 7 days of captivity. Taken together, our results indicate that lead-phase forms of P. cinereus have higher stress levels than the red-stripe forms, which may be a consequence of their higher exposure to, and/or attacks from, predators. They may also indicate that the lead-phase form is less-suited to captivity than the red-stripe form of this species [Current Zoology 56 (2: 238–243, 2010].
Hocking, Daniel J; Babbitt, Kimberly J
Ecosystems provide a vast array of services for human societies, but understanding how various organisms contribute to the functions that maintain these services remains an important ecological challenge. Predators can affect ecosystem functions through a combination of top-down trophic cascades and bottom-up effects on nutrient dynamics. As the most abundant vertebrate predator in many eastern US forests, woodland salamanders (Plethodon spp.) likely affect ecosystems functions. We examined the effects of red-backed salamanders (Plethodon cinereus) on a variety of forest ecosystem functions using a combined approach of large-scale salamander removals (314-m(2) plots) and small-scale enclosures (2 m(2)) where we explicitly manipulated salamander density (0, 0.5, 1, 2, 4 m(-2)). In these experiments, we measured the rates of litter and wood decomposition, potential nitrogen mineralization and nitrification rates, acorn germination, and foliar insect damage on red oak seedlings. Across both experimental venues, we found no significant effect of red-backed salamanders on any of the ecosystem functions. We also found no effect of salamanders on intraguild predator abundance (carabid beetles, centipedes, spiders). Our study adds to the already conflicting evidence on effects of red-backed salamander and other amphibians on terrestrial ecosystem functions. It appears likely that the impact of terrestrial amphibians on ecosystem functions is context dependent. Future research would benefit from explicitly examining terrestrial amphibian effects on ecosystem functions under a variety of environmental conditions and in different forest types.
Daniel J Hocking
Full Text Available Ecosystems provide a vast array of services for human societies, but understanding how various organisms contribute to the functions that maintain these services remains an important ecological challenge. Predators can affect ecosystem functions through a combination of top-down trophic cascades and bottom-up effects on nutrient dynamics. As the most abundant vertebrate predator in many eastern US forests, woodland salamanders (Plethodon spp. likely affect ecosystems functions. We examined the effects of red-backed salamanders (Plethodon cinereus on a variety of forest ecosystem functions using a combined approach of large-scale salamander removals (314-m(2 plots and small-scale enclosures (2 m(2 where we explicitly manipulated salamander density (0, 0.5, 1, 2, 4 m(-2. In these experiments, we measured the rates of litter and wood decomposition, potential nitrogen mineralization and nitrification rates, acorn germination, and foliar insect damage on red oak seedlings. Across both experimental venues, we found no significant effect of red-backed salamanders on any of the ecosystem functions. We also found no effect of salamanders on intraguild predator abundance (carabid beetles, centipedes, spiders. Our study adds to the already conflicting evidence on effects of red-backed salamander and other amphibians on terrestrial ecosystem functions. It appears likely that the impact of terrestrial amphibians on ecosystem functions is context dependent. Future research would benefit from explicitly examining terrestrial amphibian effects on ecosystem functions under a variety of environmental conditions and in different forest types.
Dinsmore, C E
Tail skin cuffs have been grafted to the upper forelimb of red-backed salamanders in either normal or 180 degrees-rotated dorsoventral orientation. Subsequent amputation through the graft region resulted in arrested regeneration, distally deficient or typical four-digit regenerates. Distribution was not substantially influenced by graft orientation nor were there any supernumerary limbs induced by the axially dislocated tail skin on the limb stumps. Furthermore, regenerates bore no indication of tail-like structures other than large granular skin glands proximally. These data indicate that, unlike limb skin, tail skin is not morphogenetically active in the epimorphic process of limb regeneration. In addition, the species used in this study is a fin-less, round-tailed salamander. It is therefore suggested that the previously reported morphogenetic effects of tail skin on limb regeneration may be related to the presence of tail fins on the species studied.
Jung, R.E.; Ward, W.L.; Kings, C.O.; Weir, L.A.
In 1999 at the Patuxent Research Refuge, we observed a large rove beetle (Staphylinus maculosus) consuming an eviscerated redback salamander (Plethodon cinereus) underneath a coverboard. Rove beetles typically eat invertebrates.
Gibbs, James P; Karraker, Nancy E
Several studies have reported climate-associated changes in phenotypically plastic traits of amphibians, yet it remains unknown whether amphibians can manifest an evolutionary response to global climate change at the rate and magnitude that it is occurring. To assess this issue, we examined temporal change in the morphology of the red-backed salamander (Plethodon cinereus), a small, abundant woodland salamander distributed widely in eastern North America with two distinct morphotypes: striped individuals associated with cooler microclimates and unstriped individuals associated with warmer microclimates. We compiled morph frequencies for 50,960 individual salamanders from 558 sites as recorded in the published literature and in unpublished field notes of herpetologists between 1908 and 2004. We observed that striping probability increased with increasing latitude, longitude, and elevation and decreased (from 80% to 74% range wide) with time. The combined forces of regional climate warming and, particularly, forest disturbance have evidently been sufficient to cause morphological evolution in this amphibian over the last century.
Carpenter, D.W.; Jung, R.E.; Sites, J.W.
The Shenandoah salamander (Plethodon shenandoah) is restricted to three isolated talus outcrops in Shenandoah National Park, VA, USA and has one of the smallest ranges of any tetrapod vertebrate. This species was listed as endangered under the US Endangered Species Act in 1989 over concern that direct competition with the red-backed salamander (Plethodon cinereus), successional habitat changes, and human impacts may cause its decline and possible extinction. We address two issues herein: (1) whether extensive introgression (through long-term hybridization) is present between the two species and threatens the survival of P. shenandoah, and (2) the level of population structure within P. shenandoah. We provide evidence from mtDNA haplotypes that shows no genetic differentiation among the three isolates of P. shenandoah, suggesting that their fragmentation is a geologically recent event, and/or that the isolates are still connected by occasional gene flow. There is also no evidence for extensive introgression of alleles in either direction between P. cinereus and P. shenandoah, which suggests that P. shenandoah may not be in danger of being genetically swamped out through hybridization with P. cinereus.
Uller, Claudia; Jaeger, Robert; Guidry, Gena; Martin, Carolyn
Techniques traditionally used in developmental research with infants have been widely used with nonhuman primates in the investigation of comparative cognitive abilities. Recently, researchers have shown that human infants and monkeys select the larger of two numerosities in a spontaneous forced-choice discrimination task. Here we adopt the same method to assess in a series of experiments spontaneous choice of the larger of two numerosities in a species of amphibian, red-backed salamanders ( Plethodon cinereus). The findings indicate that salamanders "go for more," just like human babies and monkeys. This rudimentary capacity is a type of numerical discrimination that is spontaneously present in this amphibian.
Hom; Jaeger; Willits
Social interactions in red-backed salamanders, Plethodon cinereusare complex and often depend upon the sex and reproductive status of individuals. In the population at Mountain Lake Biological Station, Virginia, adult males outnumber sexually active (gravid) females by approximately 2:1, because males court annually while females accept mates only biennially. Field observations of male P. cinereus suggest that males maintain feeding territories and allow non-gravid (not sexually active) females access to their territories for foraging, whereas they defend their territories against intrusion by males. We used a game-theoretic analysis of models of male fitness to examine conditions that would favour the evolution of this 'permissive' behaviour. Under a series of increasingly realistic behavioural assumptions, we found that the critical factor in the evolution of male permissive behaviour is female preference for permissive males. If non-gravid females are more likely to return to mate with a male that allows access to his territory, then permissive behaviour is likely to evolve. Furthermore, the evolution of permissive behaviour is facilitated when energetic losses due to territorial defence exceed those due to allowing females to forage.1997The Association for the Study of Animal Behaviour
Davis, Andrew K; Cecala, Kristen
Reports of an unusual intraerythrocytic pathogen in amphibian blood have been made for decades; these pathogens appear as membrane-bound vacuoles within erythrocytes. It is now understood that the pathogen is a Rickettsia bacteria, which are obligate intracellular parasites, and most are transmitted by arthropod vectors. In an effort to further understand the host range and characteristics of this pathogen, we examined 20 Ocoee salamanders (Desmognathus ocoee) from a site in southwest North Carolina for the presence of rickettsial inclusions and report the general characteristics of infections. Seven individuals (35%) were infected, and this level of prevalence was consistent with all other members of this genus examined to date. In contrast, infections within the genus Plethodon tend to occur in less than 10% of individuals, based on review of the literature. The difference could be related to the semi-aquatic nature of Desmognathus salamanders versus the completely terrestrial Plethodon. It is also possible that both groups are hosts to different types of rickettsial bacteria, since the inclusions found here only vaguely resembled those found recently in red-backed salamanders (Plethodon cinereus). Inclusions in Ocoee salamanders measured 4 microm in diameter on average, and most cells (88%) only had one inclusion. Of all infected individuals, inclusions occurred in approximately 3% of erythrocytes. Similar to that found in red-backed salamanders, infected hosts were significantly larger than non-infected. Future investigations into the nature of this unusual parasite of amphibians would help to further our understanding of its life cycle and transmission.
This work presents a refined staging table for the direct-developing red-backed salamander Plethodon cinereus, which is based on the incomplete staging system of James Norman Dent (J Morphol 1942; 71:577-601). This common species from eastern North America is a member of the species-rich lungless salamander family Plethodontidae. The staging table presented here covers several stages omitted by Dent and reveals novel developmental features of P. cinereus embryos. These include putative Leydig cells and open gill clefts, which are found in larvae of metamorphosing species but were previously reported as absent in direct-developing Plethodon. Other features found in larvae of metamorphosing salamander species, such as the palatopterygoid bone and lateral line neuromasts, were not observed in this material. The occurrence of larval and metamorphic features in these embryos has direct bearing on the patterns of life history evolution within the plethodontidae family. This study emphasizes the degree to which typically larval structures are retained in this direct-developing species and provides a staging table for further investigations into the development and evolution of plethodontid salamanders. Copyright © 2011 Wiley-Liss, Inc.
Dinsmore, C E; Maren, T H
Acetazolamide, a potent and highly specific inhibitor of carbonic anhydrase, is teratogenic in mammalian embryos and when administered during early limb development causes unique limb defects in a time- and dose-dependent manner. The regenerating urodele limb is often considered to be a good experimental analog of limb development and, if it employs the same mechanisms of tissue interactions during pattern formation, should be susceptible to teratogens which selectively disrupt developmental limb patterning. This study demonstrates that while carbonic anhydrase inhibition is toxic to the red-backed salamander, Plethodon cinereus, it does not have the same teratogenic effect on limb regeneration as seen in mammalian limb development. Several points are considered as to why the regenerating limb, at least in this salamander species, may not be suitable for studying this class of teratogen.
Nancy R.KOHN; Jennifer M.DEITLOFF; Schuyler F.DARTEZ; Michelle M.WILCOX; Robert G.JAEGER
We investigated how exposure duration (time that two individuals initially interact) and separation interval (time between the initial interaction and a subsequent interaction) affect recognition memory of conspecifics in male red-backed salamanders Plethodon cinereus.Previous studies have demonstrated that this species aggressively defends territories.We recorded aggressive behavior to assess recognition memory,because aggression is more intense toward previously unencountered individuals compared to previously encountered individuals in this species.We found that with 15-min exposures and 5-day separation intervals,focal males did not spend significantly more time threatening ‘unfamiliar' intruders than ‘familiar' intruders.After either 8-hour exposures and 5-day separation intervals and 5-day exposures and 5-day separation intervals,focal males spent significantly more time threatening unfamiliar intruders than familiar intruders.These results suggest that male red-backed salamanders can remember familiar conspecifics (e.g.,territorial neighbors) after at least an 8-hour exposure duration and that memory persists at least as long as 5 days.After 5-day exposure and 15-day separation intervals,we found no significant difference in aggressive behavior toward familiar and unfamiliar intruders.Long separation intervals (15 days) may lead either to loss of memory of previously familiar individuals or,alternatively,aggressive reassessment of individuals as only a change in behavior indicates positively that memory has occurred.Thus,variance in territorial defense within an individual may depend on its ability to recognize conspecific males.
Nancy R. KOHN, Jennifer M. DEITLOFF, Schuyler F. DARTEZ, Michelle M. WILCOX, Robert G. JAEGER
Full Text Available We investigated how exposure duration (time that two individuals initially interact and separation interval (time between the initial interaction and a subsequent interaction affect recognition memory of conspecifics in male red-backed salamanders Plethodon cinereus. Previous studies have demonstrated that this species aggressively defends territories. We recorded aggressive behavior to assess recognition memory, because aggression is more intense toward previously unencountered individuals compared to previously encountered individuals in this species. We found that with 15-min exposures and 5-day separation intervals, focal males did not spend significantly more time threatening ‘unfamiliar’ intruders than ‘familiar’ intruders. After either 8-hour exposures and 5-day separation intervals and 5-day exposures and 5-day separation intervals, focal males spent significantly more time threatening unfamiliar intruders than familiar intruders. These results suggest that male red-backed salamanders can remember familiar conspecifics (e.g., territorial neighbors after at least an 8-hour exposure duration and that memory persists at least as long as 5 days. After 5-day exposure and 15-day separation intervals, we found no significant difference in aggressive behavior toward familiar and unfamiliar intruders. Long separation intervals (15 days may lead either to loss of memory of previously familiar individuals or, alternatively, aggressive reassessment of individuals as only a change in behavior indicates positively that memory has occurred. Thus, variance in territorial defense within an individual may depend on its ability to recognize conspecific males [Current Zoology 59 (3: 326–334, 2013].
Brucker, Robert M; Baylor, Cambria M; Walters, Robert L; Lauer, Antje; Harris, Reid N; Minbiole, Kevin P C
Beneficial bacteria that live on salamander skins have the ability to inhibit pathogenic fungi. Our study aimed to identify the specific chemical agent(s) of this process and asked if any of the antifungal compounds known to operate in analogous plant-bacteria-fungi systems were present. Crude extracts of bacteria isolated from salamander skin were exposed to HPLC, UV-Vis, GC-MS, and HR-MS analyses. These investigations show that 2,4-diacetylphloroglucinol is produced by the bacteria isolate Lysobacter gummosus (AB161361), which was found on the red-backed salamander, Plethodon cinereus. Furthermore, exposure of the amphibian fungal pathogen, Batrachochytrium dendrobatidis (isolate JEL 215), to different concentrations of 2,4-diacetylphloroglucinol resulted in an IC50 value of 8.73 microM, comparable to crude extract concentrations. This study is the first to show that an epibiotic bacterium on an amphibian species produces a chemical that inhibits pathogenic fungi.
Nancy R. KOHN Jennifer M. DEITLOFF Schuyler F. DARTEZ Michelle M.WILCOX Robert G. JAEGER
...） and separation interval （time be- tween the initial interaction and a subsequent interaction） affect recognition memory of conspecifics in male red-backed sala- manders Plethodon cinereus...
Brucker, Robert M; Harris, Reid N; Schwantes, Christian R; Gallaher, Thomas N; Flaherty, Devon C; Lam, Brianna A; Minbiole, Kevin P C
Disease has spurred declines in global amphibian populations. In particular, the fungal pathogen Batrachochytrium dendrobatidis has decimated amphibian diversity in some areas unaffected by habitat loss. However, there is little evidence to explain how some amphibian species persist despite infection or even clear the pathogen beyond detection. One hypothesis is that certain bacterial symbionts on the skin of amphibians inhibit the growth of the pathogen. An antifungal strain of Janthinobacterium lividum, isolated from the skin of the red-backed salamander Plethodon cinereus, produces antifungal metabolites at concentrations lethal to B. dendrobatidis. Antifungal metabolites were identified by using reversed phase high performance liquid chromatography, high resolution mass spectrometry, nuclear magnetic resonance, and UV-Vis spectroscopy and tested for efficacy of inhibiting the pathogen. Two metabolites, indole-3-carboxaldehyde and violacein, inhibited the pathogen's growth at relatively low concentrations (68.9 and 1.82 microM, respectively). Analysis of fresh salamander skin confirmed the presence of J. lividum and its metabolites on the skin of host salamanders in concentrations high enough to hinder or kill the pathogen (51 and 207 microM, respectively). These results support the hypothesis that cutaneous, mutualistic bacteria play a role in amphibian resistance to fungal disease. Exploitation of this biological process may provide long-term resistance to B. dendrobatidis for vulnerable amphibians and serve as a model for managing future emerging diseases in wildlife populations.
Adam L. CRANE, Alicia MATHIS
Full Text Available Although salamanders have been shown to respond to classical conditioning, spatial learning has been largely unstudied. We tested whether salamanders could learn to locate foraging areas by using landmarks. We trained 10 salamanders Plethodon angusticlavius to use landmarks (small rocks to locate patches within the arena containing food (blackworms Lumbriculus variegatus. At the corners of each square testing arena were four plastic dishes, one containing blackworms and the other three empty. A rock was placed in front of the dish containing blackworms, and the location of the food-dish was randomly chosen for each training trial. A control group was also trained to feed on blackworms in the presence of a rock, but the rock was positioned randomly among the four dish locations so that the rock was not a reliable landmark for the worms. Although the length of the training period for individual salamanders varied (22–38 trainings per individual, the mean number of trainings for salamanders in the control and experimental groups was equal (30 training trials. During testing, no blackworms were present to eliminate any visual or chemical cues emanating directly from the prey. Individuals trained with the rock landmarks spent significantly more time in the area of the landmark than did control salamanders [Current Zoology 57 (4: 485–490, 2011].
Sites, J.W.; Morando, M.; Highton, R.; Huber, F.; Jung, R.E.
The Shenandoah salamander (Plethodon shenandoah), known from isolated talus slopes on three of the highest mountains in Shenandoah National Park, is listed as state-endangered in Virginia and federally endangered under the U.S. Endangered Species Act. A 1999 paper by G. R. Thurow described P. shenandoah-like salamanders from three localities further south in the Blue Ridge Physiographic Province, which, if confirmed, would represent a range extension for P. shenandoah of approximately 90 km from its nearest known locality. Samples collected from two of these three localities were included in a molecular phylogenetic study of the known populations of P. shenandoah, and all other recognized species in the Plethodon cinereus group, using a 792 bp region of the mitochondrial cytochrome-b gene. Phylogenetic estimates were based on Bayesian, maximum likelihood, and maximum parsimony methods and topologies examined for placement of the new P. shenandoah-like samples relative to all others. All topologies recovered all haplotypes of the P. shenandoah-like animals nested within P. cinereus, and a statistical comparison of the best likelihood tree topology with one with an enforced (Thurow + Shenandoah P. shenandoah) clade revealed that the unconstrained tree had a significantly lower -In L score (P Park.
Carly Muletz; Caruso, Nicholas M.; Fleischer, Robert C.; McDiarmid, Roy W.; Lips, Karen R.
Widespread population declines in terrestrial Plethodon salamanders occurred by the 1980s throughout the Appalachian Mountains, the center of global salamander diversity, with no evident recovery. We tested the hypothesis that the historic introduction and spread of the pathogenic fungus Batrachochytrium dendrobatidis (Bd) into the eastern US was followed by Plethodon population declines. We expected to detect elevated prevalence of Bd prior to population declines as observed for Central Amer...
Cabe, P R; Page, R B; Hanlon, T J; Aldrich, M E; Connors, L; Marsh, D M
Several recent studies have shown that amphibian populations may exhibit high genetic subdivision in areas with recent fragmentation and urban development. Less is known about the potential for genetic differentiation in continuous habitats. We studied genetic differentiation of red-backed salamanders (Plethodon cinereus) across a 2-km transect through continuous forest in Virginia, USA. Mark-recapture studies suggest very little dispersal for this species, whereas homing experiments and post-Pleistocene range expansion both suggest greater dispersal abilities. We used six microsatellite loci to examine genetic population structure and differentiation between eight subpopulations of red-backed salamanders at distances from 200 m to 2 km. We also used several methods to extrapolate dispersal frequencies and test for sex-biased dispersal. We found small, but detectable differentiation among populations, even at distances as small as 200 m. Differentiation was closely correlated with distance and both Mantel tests and assignment tests were consistent with an isolation-by-distance model for the population. Extrapolations of intergenerational variance in spatial position (sigma(2)<15 m(2)) and pair-wise dispersal frequencies (4 Nm < 25 for plots separated by 300 m) both suggest limited gene flow. Additionally, tests for sex-biased dispersal imply that dispersal frequency is similarly low for both sexes. We suggest that these low levels of gene flow and the infrequent dispersal observed in mark-recapture studies may be reconciled with homing ability and range expansion if dispersing animals rarely succeed in breeding in saturated habitats, if dispersal is flexible depending on the availability of habitat, or if dispersal frequency varies across the geographic range of red-backed salamanders.
Kerney, Ryan R; Blackburn, David C; Müller, Hendrik; Hanken, James
Recent molecular phylogenies suggest the surprising reacquisition of posthatching metamorphosis within an otherwise direct-developing clade of lungless salamanders (family Plethodontidae). Metamorphosis was long regarded as plesiomorphic for plethodontids, yet the genus Desmognathus, which primarily includes metamorphosing species, is now nested within a much larger clade of direct-developing species. The extent to which the putative reacquisition of metamorphosis in Desmognathus represents a true evolutionary reversal is contingent upon the extent to which both larva-specific features and metamorphosis were actually lost during the evolution of direct development. In this study we analyze development of the hyobranchial skeleton, which is dramatically remodeled during salamander metamorphosis, in the direct-developing red-backed salamander, Plethodon cinereus. We find dramatic remodeling of the hyobranchial skeleton during embryogenesis in P. cinereus and the transient appearance of larva-specific cartilages. Hyobranchial development in this direct-developing plethodontid is highly similar to that in metamorphosing plethodontids (e.g., Desmognathus). The proposed reacquisition of hyobranchial metamorphosis within Desmognathus does not represent the "re-evolution" of a lost phenotype, but instead the elaboration of an existing developmental sequence. © 2011 The Author(s). Evolution © 2011 The Society for the Study of Evolution.
Full Text Available Widespread population declines in terrestrial Plethodon salamanders occurred by the 1980s throughout the Appalachian Mountains, the center of global salamander diversity, with no evident recovery. We tested the hypothesis that the historic introduction and spread of the pathogenic fungus Batrachochytrium dendrobatidis (Bd into the eastern US was followed by Plethodon population declines. We expected to detect elevated prevalence of Bd prior to population declines as observed for Central American plethodontids. We tested 1,498 Plethodon salamanders of 12 species (892 museum specimens, 606 wild individuals for the presence of Bd, and tested 94 of those for Batrachochytrium salamandrivorans (Bs and for ranavirus. Field samples were collected in 2011 from 48 field sites across a 767 km transect. Historic samples from museum specimens were collected at five sites with the greatest number and longest duration of collection (1957-987, four of which were sampled in the field in 2011. None of the museum specimens were positive for Bd, but four P. cinereus from field surveys were positive. The overall Bd prevalence from 1957-2011 for 12 Plethodon species sampled across a 757 km transect was 0.2% (95% CI 0.1-0.7%. All 94 samples were negative for Bs and ranavirus. We conclude that known amphibian pathogens are unlikely causes for declines in these Plethodon populations. Furthermore, these exceptionally low levels of Bd, in a region known to harbor Bd, may indicate that Plethodon specific traits limit Bd infection.
Muletz, Carly; Caruso, Nicholas M; Fleischer, Robert C; McDiarmid, Roy W; Lips, Karen R
Widespread population declines in terrestrial Plethodon salamanders occurred by the 1980s throughout the Appalachian Mountains, the center of global salamander diversity, with no evident recovery. We tested the hypothesis that the historic introduction and spread of the pathogenic fungus Batrachochytrium dendrobatidis (Bd) into the eastern US was followed by Plethodon population declines. We expected to detect elevated prevalence of Bd prior to population declines as observed for Central American plethodontids. We tested 1,498 Plethodon salamanders of 12 species (892 museum specimens, 606 wild individuals) for the presence of Bd, and tested 94 of those for Batrachochytrium salamandrivorans (Bs) and for ranavirus. Field samples were collected in 2011 from 48 field sites across a 767 km transect. Historic samples from museum specimens were collected at five sites with the greatest number and longest duration of collection (1957-987), four of which were sampled in the field in 2011. None of the museum specimens were positive for Bd, but four P. cinereus from field surveys were positive. The overall Bd prevalence from 1957-2011 for 12 Plethodon species sampled across a 757 km transect was 0.2% (95% CI 0.1-0.7%). All 94 samples were negative for Bs and ranavirus. We conclude that known amphibian pathogens are unlikely causes for declines in these Plethodon populations. Furthermore, these exceptionally low levels of Bd, in a region known to harbor Bd, may indicate that Plethodon specific traits limit Bd infection.
Muletz, Carly; Caruso, Nicholas M.; Fleischer, Robert C.; McDiarmid, Roy W.; Lips, Karen R.
Widespread population declines in terrestrial Plethodon salamanders occurred by the 1980s throughout the Appalachian Mountains, the center of global salamander diversity, with no evident recovery. We tested the hypothesis that the historic introduction and spread of the pathogenic fungus Batrachochytrium dendrobatidis (Bd) into the eastern US was followed by Plethodon population declines. We expected to detect elevated prevalence of Bd prior to population declines as observed for Central American plethodontids. We tested 1,498 Plethodon salamanders of 12 species (892 museum specimens, 606 wild individuals) for the presence of Bd, and tested 94 of those for Batrachochytrium salamandrivorans (Bs) and for ranavirus. Field samples were collected in 2011 from 48 field sites across a 767 km transect. Historic samples from museum specimens were collected at five sites with the greatest number and longest duration of collection (1957–987), four of which were sampled in the field in 2011. None of the museum specimens were positive for Bd, but four P. cinereus from field surveys were positive. The overall Bd prevalence from 1957–2011 for 12 Plethodon species sampled across a 757 km transect was 0.2% (95% CI 0.1–0.7%). All 94 samples were negative for Bs and ranavirus. We conclude that known amphibian pathogens are unlikely causes for declines in these Plethodon populations. Furthermore, these exceptionally low levels of Bd, in a region known to harbor Bd, may indicate that Plethodon specific traits limit Bd infection. PMID:25084159
Ransom, Tami S
In addition to creating or modifying habitat, ecosystem engineers interact with other species as predators, prey, or competitors. The earthworm, Lumbricus terrestris, interacts with the common woodland salamander, Plethodon cinereus, via: (1) ecosystem engineering, by providing burrows that are used as a refuge, (2) direct effects as a prey item, and (3) indirectly, by competing with microinvertebrates, another prey item for P. cinereus. Using enclosures in the forest, I examined the relative strengths of these component pathways between seasons and salamander age classes. I found that the relative strength (partial eta2) of the positive direct (trophic) effect of L. terrestris on the change in mass of P. cineresus was greater than that of the negative indirect effect, but only in summer. Positive effects of ecosystem engineering were only evident over the winter as increased adult survival. This research has implications for how habitat provisioning complements more well-studied species interactions, such as competition and predation, within communities.
Muzzall, P M; Peebles, C R; Burton, T M
Totals of 52 dusky salamanders Desmognathus fuscus, 51 two-lined salamanders Eurycea bislineata, 54 red-backed salamanders Plethodon cinereus, and 3 spring salamanders Gyrinophilus porphyriticus (Plethodontidae) collected in June and August 1995 from Paradise Brook, a tributary to Hubbard Brook, New Hampshire, were examined for parasites. Parasites found were Brachycoelium storeriae, Brachycoelium sp., Bothriocephalus rarus, Falcaustra sp., Omeia sp., Batracholandros magnavulvaris, and Cepedietta michiganensis. Eighty-six percent of the red-backed salamanders, a terrestrial species, harbored 1 or more parasites. Among the aquatic and semiaquatic species, 27% of the dusky and 45% of the two-lined salamanders were infected with 1 or more parasites.
Morgan, G T
Tritiated-thymidine autoradiography was used to follow the progress of cells through meiosis in male P. cinereus at 20 degrees C. Spermatocytes spend 7 days in leptotene, 5 days in zygotene and 3 days in pachytene before entering the diffuse stage. Diffuse lasts for 8 days and is followed by a diplotene of 2 days. First and second meiotic metaphase occur a total of 26 days after the end of premeiotic S. Considering the information for P. cinereus together with that for 3 other species, it appears that in amphibians the duration of meiosis does not show the linear relation with C-value that has been described for other groups of organisms.
DeGross, Douglas J.; Bury, R. Bruce
The Plethodon elongatus Complex in the Klamath-Siskiyou Ecoregion of southern Oregon and northern California includes three species: the Del Norte salamander, Plethodon elongatus; the Siskiyou Mountains salamander, P. stormi; and the Scott Bar salamander, P. asupak. This review aims to summarize the current literature and information available on select topics for P. stormi and P. asupak. These are both terrestrial salamanders belonging to the Family Plethodontidae, which contains more species and has a wider geographic distribution than any other family of salamanders (Wake 1966, 2006; Pough 1989). The genera of this family have greatly diversified ecologically across North America, Central America, northern South America, Sardinia, southeastern France and northwestern Italy, and have recently been discovered on the Korean peninsula (Min et al. 2005). The genus Plethodon is found exclusively in North America and is split into three distinct clades, based upon morphology and phylogenetics (Highton and Larson 1979): eastern small Plethodon, eastern large Plethodon, and the western Plethodon. The western Plethodon are the greatest representation of Plethodontidae in the Pacific Northwest, with 8 species. The two species with the most restricted ranges of these regional congeners are the Siskiyou Mountains and Scott Bar salamanders. These salamanders occupy the interior of the Klamath-Siskiyou Ecoregion which straddles the California and Oregon state lines, between Siskiyou County (CA) and Jackson and Josephine Counties (OR). The relatively recent discovery of P. asupak (Mead et al. 2005) and the limited range of both species have created an environment of uncertain conservation status for these species. This review will focus on four central topics of concern for land and resource managers: Biology; Taxonomy; Habitat; and Detection Probabilities/Occupancy.
Wilburn, Damien B; Bowen, Kathleen E; Feldhoff, Pamela W; Feldhoff, Richard C
The evolutionary success of plethodontid salamanders for ~100 MY is due partly to the use of courtship pheromones that regulate female receptivity. In ~90 % of plethodontid species, males deliver pheromones by "scratching" a female's dorsum, where pheromones diffuse transdermally into the bloodstream. However, in a single clade, representing ~10 % of Plethodon spp., males apply pheromones to the female's nares for olfactory delivery. Molecular studies have identified three major pheromone families: Plethodontid Receptivity Factor (PRF), Plethodontid Modulating Factor (PMF), and Sodefrin Precursor-like Factor (SPF). SPF and PMF genes are relatively ancient and found in all plethodontid species; however, PRF is found exclusively in the genus Plethodon - which includes species with transdermal, olfactory, and intermediate delivery behaviors. While previous proteomic analyses suggested PRF and PMF are dominant in slapping species and SPF is dominant in non-Plethodon scratching species, it was unclear how protein expression of different pheromone components may vary across delivery modes within Plethodon. Therefore, the aim of this study was to proteomically characterize the pheromones of a key scratching species in this evolutionary transition, Plethodon cinereus. Using mass spectrometry-based techniques, our data support the functional replacement of SPF by PRF in Plethodon spp. and an increase in PMF gene duplication events in both lineage-dependent and delivery-dependent manners. Novel glycosylation was observed on P. cinereus PRFs, which may modulate the metabolism and/or mechanism of action for PRF in scratching species. Cumulatively, these molecular data suggest that the replacement of pheromone components (e.g., SPF by PRF) preceded the evolutionary transition of the functional complex from transdermal to olfactory delivery.
Pelletier, Tara A; Crisafulli, Charlie; Wagner, Steve; Zellmer, Amanda J; Carstens, Bryan C
Allopatry is commonly used to predict boundaries in species delimitation investigations under the assumption that currently allopatric distributions are indicative of reproductive isolation; however, species ranges are known to change over time. Incorporating a temporal perspective of geographic distributions should improve species delimitation; to explore this, we investigate three species of western Plethodon salamanders that have shifted their ranges since the end of the Pleistocene. We generate species distribution models (SDM) of the current range, hindcast these models onto a climatic model 21 Ka, and use three molecular approaches to delimit species in an integrated fashion. In contrast to expectations based on the current distribution, we detect no independent lineages in species with allopatric and patchy distributions (Plethodon vandykei and Plethodon larselli). The SDMs indicate that probable habitat is more expansive than their current range, especially during the last glacial maximum (LGM) (21 Ka). However, with a contiguous distribution, two independent lineages were detected in Plethodon idahoensis, possibly due to isolation in multiple glacial refugia. Results indicate that historical SDMs are a better predictor of species boundaries than current distributions, and strongly imply that researchers should incorporate SDM and hindcasting into their investigations and the development of species hypotheses. © The Author(s) 2014. Published by Oxford University Press, on behalf of the Society of Systematic Biologists. All rights reserved. For Permissions, please email: email@example.com.
Wyman, R.L.; Hawksley-Lescault, D.S.
Censuses at two sites in Delaware County, New York from spring 1981 through spring 1985 indicated that the density and distribution of Plethodon cinereus were influenced by soil pH but not by soil temperature or moisture. Of 1044 1-m/sup 2/ quadrats of forest litter searched, 284 had a pH of 3.7 or less and only 25 of these (8.8%) contained salamanders. Of 760 quadrats with a pH 3.8 or more, 386 (50.8%) contained salamanders. Juvenile salamanders were never found on soils with a pH less than or equal to 3.7. Seasonal salamander density was correlated (r = -0.92) with the percentage of quadrats with a pH of 3.7 and less. Salamanders apparently were excluded from 27% of forest habitat because of low soil pH. In the laboratory, P. cinereus preferred to occupy substrates near neutral pH when given a choice among three levels of substrate acidity. The acutely lethal pH was between 2.5 and 3 and the 8-mo chronically lethal pH was between 3 and 4. Growth and respiration were reduced at low pHs. The influence of soil pH on salamander distribution might fundamentally change the forest floor decomposer food web of which P. cinereus is an upper-level consumer.
Dinsmore, C E
Regeneration of the tail in the plethodontid salamander, Plethodon cinereus, occurs following either surgical amputation or induced autotomy. Autotomy may occur along any one of the caudal myosepta which form natural cleavage planes. The distally attached myofibers break away from the myoseptum which then becomes part of the stump surface remaining intact during subsequent regeneration of the distal segments. Under these conditions, therefore, muscle fibers do not normally participate in tail regeneration. If, however, the myofibers of a caudal myotome are interrupted as in mid-segment amputation, the damaged fibers undergo hyaline degeneration and are largely replaced by connective tissue often as far proximally as the next myoseptum. Other tissues at the amputation surface are concurrently engaged in the epimorphic regeneration of the tail, including myogenesis, though at a slower initial rate than that following autotomy. Muscle cells, therefore, appear not to participate in epimorphic tail regeneration in Plethodon cinereus following either surgical amputation or induced autotomy.
Johnson, M S; Paulus, H I; Salice, C J; Checkai, R T; Simini, M
Red-backed salamanders (Plethodon cinereus) were exposed to four different concentrations of 1,3,5-trinitrohexahydro-1,3,5-triazine (RDX) in soil under controlled laboratory conditions for 28 days. Wild-caught P. cinereus (N = 20/treatment) were exposed to target concentrations of 5,000, 1000, 100, 10, and 0 mg RDX/kg soil (dry wt) using a microcosm design. Animals were fed 5 to 10 uncontaminated mutant Drosophila flies every 3 days and monitored daily. Animals were weighed 1 day before being placed in treatment and weekly thereafter. RDX concentrations in soil were analytically determined after the compound was added and mixed at the beginning, the midpoint, and the end of exposure. RDX soil concentrations were relatively stable throughout the exposure period. Signs of overt toxicity were observed primarily in the highest exposure group. Salamanders exposed to 5,000 mg RDX/kg soil exhibited signs of neuromuscular effects (lethargy, gaping, hypersensitivity, tremors) and exhibited significant weight loss. A single moribund animal from this group lost >20% of its original body weight and was killed. Animals in this exposure group also lost weight relative to animals in other treatments. Histopathologic evaluations, including an evaluation of melanomacrophage parameters, indicated no strong treatment-related findings. This study provided information regarding the effects from subchronic dermal exposure of a terrestrial amphibian species to RDX in soil and provides a microcosm approach to the evaluation of toxicity of contaminants in soil to a terrestrial vertebrate.
Feldhoff Pamela W
Full Text Available Abstract Background The mental gland pheromone of male Plethodon salamanders contains two main protein components: a 22 kDa protein named Plethodon Receptivity Factor (PRF and a 7 kDa protein named Plethodon Modulating Factor (PMF, respectively. Each protein component individually has opposing effects on female courtship behavior, with PRF shortening and PMF lengthening courtship. In this study, we test the hypothesis that PRF or PMF individually activate vomeronasal neurons. The agmatine-uptake technique was used to visualize chemosensory neurons that were activated by each protein component individually. Results Vomeronasal neurons exposed to agmatine in saline did not demonstrate significant labeling. However, a population of vomeronasal neurons was labeled following exposure to either PRF or PMF. When expressed as a percent of control level labeled cells, PRF labeled more neurons than did PMF. These percentages for PRF and PMF, added together, parallel the percentage of labeled vomeronasal neurons when females are exposed to the whole pheromone. Conclusion This study suggests that two specific populations of female vomeronasal neurons are responsible for responding to each of the two components of the male pheromone mixture. These two neural populations, therefore, could express different receptors which, in turn, transmit different information to the brain, thus accounting for the different female behavior elicited by each pheromone component.
US Fish and Wildlife Service, Department of the Interior — The Cheat Mountain Salamander (Plethodon nettingi) is an endemic species native only to West Virginia. Besides being restricted to one state, Cheat Mountain...
US Fish and Wildlife Service, Department of the Interior — The Cheat Mountain salamander (CMS; Plethodon nettingi) is endemic to high-elevation forests of the Allegheny Mountains in Tucker, Randolph, Pocahontas, Grant, and...
Fredericks, L P; Dankert, J R
As resistance increases against fungal antibiotics, antimicrobial peptides are receiving attention as possible replacements. The dermal glands of frogs secrete, among other things, antimicrobial peptides. As part of the innate immune system, stressors may affect the production of antimicrobial peptides by dermal glands. The dermal secretions of some salamanders have been examined for their toxic secretions, but little attention has been given to salamander antimicrobial peptides. This study examines the skin from the tail region for the production of antimicrobial peptides in the terrestrial salamander, Plethodon cinereus. Fractions of tail extracts were isolated using cation-exchange chromatography and reverse-phase HPLC. An HPLC fraction eluting at 15.75 min (HPLC run: 30 min, 30-80% acetonitrile/water gradient, Aquapore RP-300 C18 column) showed activity against Staphylococcus aureus but not against Escherichia coli. The antibacterial activity gradually increased over a 4-hr incubation time up to about 85% inhibition of bacterial growth. Lysis of guinea pig red blood cells also increased gradually over a 1-hr time period. J. Exp. Zool. 287:340-345, 2000. Copyright 2000 Wiley-Liss, Inc.
Niewiarowski Peter H; Petruzzi Erin E; Moore Francisco
Abstract Background In eastern North America two common colour morphs exist in most populations of redback salamanders (Plethodon cinereus). Previous studies have indicated that the different morphs may be adapted to different thermal niches and the morphological variation has been linked to standard metabolic rate at 15°C in one population of P. cinereus. It has therefore been hypothesized that a correlated response to selection on metabolic rate across thermal niches maintains the colour po...
Crump, M L
Mean values of energetic parameters are used in ecosystem studies because only crude estimates are possible when dealing with a large scope. However, when the focus is at the population level, finer resolution is possible and thus potential ontogenetic differences should be examined. Analysis of energetic parameters of a population of terrestrial salamander, Plethodon cinereus, was undertaken in order to determine ontogenetic differences.Digestive efficiency does not differ significantly between age classes, nor is it correlated with energy intake. Adult males and non-gravid females have virtually identical mean efficiencies. Thus it is probably meaningful to calculate energy flow through the population using a mean digestive efficiency value. Percent water content decreases significantly with increasing age whereas percent ash content increases significantly. Thus when estimating energy tied up in standing crop, correction must be applied for ontogenetic differences in water and ash content. Once these corrections have been made, the AFDW-specific energy content is not significantly different between age groups (with the exception of gravid females which is higher). Salamanders accumulate energy in their tissues with increasing age, although immatures are not significantly different from non-gravid adults. Since gravid females have significantly more joules/mg wet weight than other groups, separate energy density values must be calculated for these individuals if a meaningful standing crop energy estimate is to be obtained from wet weight biomass.
Siddig, Ahmed A.; Ellison, Aaron M.; Scott Jackson
Herpetologists and conservation biologists frequently use convenient and cost-effective, but less accurate, abundance indices (e.g., number of individuals collected under artificial cover boards or during natural objects surveys) in lieu of more accurate, but costly and destructive, population size estimators to detect and monitor size, state, and trends of amphibian populations. Although there are advantages and disadvantages to each approach, reliable use of abundance indices requires that ...
Wagner, R.S.; Miller, Mark P.; Crisafulli, Charles; Haig, Susan M.
The Larch Mountain salamander (Plethodon larselli Burns, 1954) is an endemic species in the Pacific northwestern United States facing threats related to habitat destruction. To facilitate development of conservation strategies, we used DNA sequences and RAPDs (random amplified polymorphic DNA) to examine differences among populations of this species. Phylogenetic analyses of cytochrome b revealed a clade of haplotypes from populations north of the Columbia River derived from a clade containing haplotypes from the river's southwestern region. Haplotypes from southeastern populations formed a separate clade. Nucleotide diversity was reduced in northern populations relative to southern populations. These results were corroborated by analyses of RAPD loci, which revealed similar patterns of clustering and diversity. Network analyses suggested that northern populations were colonized following a range expansion mediated by individuals from populations located southwest of the river. Changes in the Columbia River's location during the Pliocene and Pleistocene likely released distributional constraints on this species, permitting their northern range expansion. Based on the barrier presented by the Columbia River's present location and differences in haplotype diversity and population structure observed between northern and southern populations, we suggest that designation of separate management units encompassing each region may assist with mitigating different threats to this species.
Liebgold, Eric B; Brodie, Edmund D; Cabe, Paul R
The local resource competition hypothesis and the local mate competition hypothesis were developed based on avian and mammalian systems to explain sex-biased dispersal. Most avian species show a female bias in dispersal, ostensibly due to resource defence, and most mammals show a male bias, ostensibly due to male-male competition. These findings confound phylogeny with mating strategy; little is known about sex-biased dispersal in other taxa. Resource defence and male-male competition are both intense in Plethodon cinereus, a direct-developing salamander, so we tested whether sex-biased dispersal in this amphibian is consistent with the local resource competition hypothesis (female-biased) or the local mate competition hypothesis (male-biased). Using fine-scale genetic spatial autocorrelation analyses, we found that females were philopatric, showing significant positive genetic structure in the shortest distance classes, with stronger patterns apparent when only territorial females were tested. Males showed no spatial genetic structure over the shortest distances. Mark-recapture observations of P. cinereus over 5 years were consistent with the genetic data: males dispersed farther than females during natal dispersal and 44% of females were recaptured within 1 m of their juvenile locations. We conclude that, in this population of a direct-developing amphibian, females are philopatric and dispersal is male-biased, consistent with the local mate competition hypothesis. © 2010 Blackwell Publishing Ltd.
Budzik, Karolina A; Żuwała, Krystyna; Kerney, Ryan
The latest research on direct developing caecilian and anuran species indicate presence of only one generation of taste organs during their ontogeny. This is distinct from indirect developing batrachians studied thus far, which possess taste buds in larvae and anatomically distinct taste discs in metamorphs. This study is a description of the tongue and taste organ morphology and development in direct developing salamander Plethodon cinereus (Plethodontidae) using histology and electron microscopy techniques. The results reveal two distinct stages tongue morphology (primary and secondary), similar to metamorphic urodeles, although only one stage of taste organ morphology. Taste disc sensory zones emerge on the surface of the oropharyngeal epithelium by the end of embryonic development, which coincides with maturation of the soft tongue. Taste organs occur in the epithelium of the tongue pad (where they are situated on the dermal papillae), the palate and the inner surface of the mandible and the maxilla. Plethodon cinereus embryos only possess taste disc type taste organs. Similar to the direct developing anuran Eleutherodactylus coqui (Eleutherodactylidae), these salamanders do not recapitulate larval taste bud morphology as an embryo. The lack of taste bud formation is probably a broadly distributed feature characteristic to direct developing batrachians. J. Morphol. 277:906-915, 2016. © 2016 Wiley Periodicals, Inc. © 2016 Wiley Periodicals, Inc.
Adams, Dean C; West, Mary E; Collyer, Michael L
1. The competitive interactions of closely related species have long been considered important determinants of community composition and a major cause of phenotypic diversification. However, while patterns such as character displacement are well documented, less is known about how local adaptation influences diversifying selection from interspecific competition. 2. We examined body size and head shape variation among allopatric and sympatric populations of two salamander species, the widespread Plethodon cinereus and the geographically restricted P. nettingi. We quantified morphology from 724 individuals from 20 geographical localities throughout the range of P. nettingi. 3. Plethodon nettingi was more robust in cranial morphology relative to P. cinereus, and sympatric localities were more robust relative to allopatric localities. Additionally, there was significantly greater sympatric head shape divergence between species relative to allopatric communities, and sympatric localities of P. cinereus exhibited greater morphological variation than sympatric P. nettingi. 4. The sympatric morphological divergence and increase in cranial robustness of one species (P. nettingi) were similar to observations in other Plethodon communities, and were consistent with the hypothesis of interspecific competition. These findings suggest that interspecific competition in Plethodon may play an important role in phenotypic diversification in this group. 5. The increase in among-population variance in sympatric P. cinereus suggests a species-specific response to divergent natural selection that is influenced in part by other factors. We hypothesize that enhanced morphological flexibility and ecological tolerance allow P. cinereus to more rapidly adapt to local environmental conditions, and initial differences among populations have allowed the evolutionary response of P. cinereus to vary across replicate sympatric locations, resulting in distinct evolutionary trajectories of
Frisbie, M.P.; Wyman, R.L. (Eastern Kentucky Univ., Richmond, KY (United States))
Earlier laboratory studies demonstrated that red-backed salamanders, Plethodon cinereus, are susceptible to osmotic disruption by low pH substrates. In natural systems, however, acidic input from precipitation may be mediated by soils before it impacts salamanders. We tested the effect of acidic rain on sodium balance in salamanders by confining individuals in enclosure in two forest types (hemlock, beech) for 34 d. Enclosures received artificial rain of either pH 3 or 5 every 3-4 d. Soils inside enclosures in the hemlock forest were more acidic than those in the beech forest at the outset. At termination, [H[sup +
MOSES, M J
An investigation of the structure of meiotic chromosomes from primary spermatocytes of two salamanders, Plethodon cinereus and Desmognathus fusca, has been made using correlated light and electron microscopy. Feulgen squashes were compared with stained sections and these related to adjacent thin sections in the electron microscope. A transition from the familiar cytological preparation to the electron image was thus effected. A linear complex consisting of three parallel strands has been observed with the electron microscope, passing along the central axis of primary spermatocyte chromosomes. The complex is similar to that found in comparable chromosomes from at least a dozen animal species. The structure in Plethodon is described in detail. Synapsis has been positively identified as the stage of meiotic prophase at which the complex occurs. Thus the complex is a part of bivalent chromosomes. It has not been seen in other stages or other divisions and is thus thought to be exclusively of synaptic occurrence. The term synaptinemal complex is suggested for the entire structure. By virtue of the material condensed around it, the complex is also seen in the light microscope where it appears as a fine, densely Feulgen-positive central core along the chromosome. The complex is thus closely associated with DNA, if not at least in part, composed of it. In the stages studied, homologous chromosomes are not always completely paired. The lateral elements of the complex separate and follow the single chromosome axes at these points. The central element disappears and thus may be a phenomenon of pairing. It is concluded that the lateral elements of the synaptinemal complex may more correctly be a "core" of the single meiotic prophase chromosome, possibly being concerned with its linear organization.
US Fish and Wildlife Service, Department of the Interior — There are two primary goals for this project: to establish baseline information on populations of the Cheat Mountain salamander and to determine if there are any...
Ziemba, Julie L; Hickerson, Cari-Ann M; Anthony, Carl D
Asian pheretimoid earthworms (e.g. Amynthas and Metaphire spp.) are invading North American forests and consuming the vital detrital layer that forest floor biota [including the keystone species Plethodon cinereus (Eastern Red-backed Salamander)], rely on for protection, food, and habitat. Plethodon cinereus population declines have been associated with leaf litter loss following the invasion of several exotic earthworm species, but there have been few studies on the specific interactions between pheretimoid earthworms and P. cinereus. Since some species of large and active pheretimoids spatially overlap with salamanders beneath natural cover objects and in detritus, they may distinctively compound the negative consequences of earthworm-mediated resource degradation by physically disturbing important salamander activities (foraging, mating, and egg brooding). We predicted that earthworms would exclude salamanders from high quality microhabitat, reduce foraging efficiency, and negatively affect salamander fitness. In laboratory trials, salamanders used lower quality microhabitat and consumed fewer flies in the presence of earthworms. In a natural field experiment, conducted on salamander populations from "non-invaded" and "pheretimoid invaded" sites in Ohio, salamanders and earthworms shared cover objects ~60% less than expected. Earthworm abundance was negatively associated with juvenile and male salamander abundance, but had no relationship with female salamander abundance. There was no effect of pheretimoid invasion on salamander body condition. Juvenile and non-resident male salamanders do not hold stable territories centered beneath cover objects such as rocks or logs, which results in reduced access to prey, greater risk of desiccation, and dispersal pressure. Habitat degradation and physical exclusion of salamanders from cover objects may hinder juvenile and male salamander performance, ultimately reducing recruitment and salamander abundance following Asian
Julie L Ziemba
Full Text Available Asian pheretimoid earthworms (e.g. Amynthas and Metaphire spp. are invading North American forests and consuming the vital detrital layer that forest floor biota [including the keystone species Plethodon cinereus (Eastern Red-backed Salamander], rely on for protection, food, and habitat. Plethodon cinereus population declines have been associated with leaf litter loss following the invasion of several exotic earthworm species, but there have been few studies on the specific interactions between pheretimoid earthworms and P. cinereus. Since some species of large and active pheretimoids spatially overlap with salamanders beneath natural cover objects and in detritus, they may distinctively compound the negative consequences of earthworm-mediated resource degradation by physically disturbing important salamander activities (foraging, mating, and egg brooding. We predicted that earthworms would exclude salamanders from high quality microhabitat, reduce foraging efficiency, and negatively affect salamander fitness. In laboratory trials, salamanders used lower quality microhabitat and consumed fewer flies in the presence of earthworms. In a natural field experiment, conducted on salamander populations from "non-invaded" and "pheretimoid invaded" sites in Ohio, salamanders and earthworms shared cover objects ~60% less than expected. Earthworm abundance was negatively associated with juvenile and male salamander abundance, but had no relationship with female salamander abundance. There was no effect of pheretimoid invasion on salamander body condition. Juvenile and non-resident male salamanders do not hold stable territories centered beneath cover objects such as rocks or logs, which results in reduced access to prey, greater risk of desiccation, and dispersal pressure. Habitat degradation and physical exclusion of salamanders from cover objects may hinder juvenile and male salamander performance, ultimately reducing recruitment and salamander abundance
I used a mark-recapture study to estimate home areas for 107 red-backed salamanders (Plethodon cinereus) in a natural forest habitat. Both males and females of this species defend feeding territories, but I presume that some individuals in this relatively highdensity population (approximately 2.8 salamanders per m(2)) are nonterritorial floaters. Although territorial salamanders exhibited greater numbers of tail autotomies, they had significantly longer relative tail lengths. This difference suggests that territorial individuals gain benefits from territorial ownership. From the observation that home area size was inversely correlated with body size, I infer that larger animals gained higher quality foraging areas. Home areas of adults were significantly more segregated intrasexually and more aggregated intersexually than would be expected from a random distribution. Furthermore, intersexual overlap of home areas was significantly greater than intrasexual home area overlap. Territorial defense of feeding areas by male and female red-backed salamanders therefore also may play a role in mating behavior.
Niewiarowski Peter H
Full Text Available Abstract Background In eastern North America two common colour morphs exist in most populations of redback salamanders (Plethodon cinereus. Previous studies have indicated that the different morphs may be adapted to different thermal niches and the morphological variation has been linked to standard metabolic rate at 15°C in one population of P. cinereus. It has therefore been hypothesized that a correlated response to selection on metabolic rate across thermal niches maintains the colour polymorphism in P. cinereus. This study tests that hypothesis. Results We found that the two colour morphs do sometimes differ in their maintenance metabolic rate (MMR profiles, but that the pattern is not consistent across populations or seasons. We also found that when MMR profiles differ between morphs those differences do not indicate that distinct niches exist. Field censuses showed that the two colour morphs are sometimes found at different substrate temperatures and that this difference is also dependent on census location and season. Conclusion While these morphs sometimes differ in their maintenance energy expenditures, the differences in MMR profile in this study are not consistent with maintenance of the polymorphism via a simple correlated response to selection across multiple niches. When present, differences in MMR profile do not indicate the existence of multiple thermal niches that consistently mirror colour polymorphism. We suggest that while a relationship between colour morph and thermal niche selection appears to exist it is neither simple nor consistent.
Petruzzi, Erin E; Niewiarowski, Peter H; Moore, Francisco B-G
In eastern North America two common colour morphs exist in most populations of redback salamanders (Plethodon cinereus). Previous studies have indicated that the different morphs may be adapted to different thermal niches and the morphological variation has been linked to standard metabolic rate at 15 degrees C in one population of P. cinereus. It has therefore been hypothesized that a correlated response to selection on metabolic rate across thermal niches maintains the colour polymorphism in P. cinereus. This study tests that hypothesis. We found that the two colour morphs do sometimes differ in their maintenance metabolic rate (MMR) profiles, but that the pattern is not consistent across populations or seasons. We also found that when MMR profiles differ between morphs those differences do not indicate that distinct niches exist. Field censuses showed that the two colour morphs are sometimes found at different substrate temperatures and that this difference is also dependent on census location and season. While these morphs sometimes differ in their maintenance energy expenditures, the differences in MMR profile in this study are not consistent with maintenance of the polymorphism via a simple correlated response to selection across multiple niches. When present, differences in MMR profile do not indicate the existence of multiple thermal niches that consistently mirror colour polymorphism. We suggest that while a relationship between colour morph and thermal niche selection appears to exist it is neither simple nor consistent.
Connette, Grant M; Semlitsch, Raymond D
Animal movement is critical to the maintenance of functional connectivity at the landscape scale and can play a key role in population persistence and metapopulation dynamics. The permeability of habitat to animal movement may vary as a result of either differential mortality, physical resistance, or simply the behavioral responses of organisms to perceived habitat quality. Understanding how and when animal movement behavior varies among habitat types is critical for identifying barriers to dispersal and predicting species distributions in relation to landscape features. We conducted an experimental translocation study and compared the movement success and behavioral strategies of plethodontid salamanders in both forest and open-canopy habitat. We found that individuals in closed-canopy forest oriented more strongly towards their home ranges and moved significantly farther on their release night. In spite of the clear differences in movement paths, the ultimate movement success of homing salamanders did not appear to vary with habitat type. Our study contributes to a growing body of literature suggesting the importance of recognizing the context dependence of animal movement behavior. Because the movement rates of displaced salamanders were significantly reduced in open-canopy, dispersal rates of plethodontid salamanders in open-canopy habitat are likely lower than in control forest. Further mechanistic studies focusing on habitat-specific movement behavior and survival costs will be valuable for effectively identifying and mitigating barriers to animal movement.
Highton, Richard; Hastings, Amy Picard; Palmer, Catherine; Watts, Richard; Hass, Carla A; Culver, Melanie; Arnold, Stevan J
Salamanders of the North American plethodontid genus Plethodon are important model organisms in a variety of studies that depend on a phylogenetic framework (e.g., chemical communication, ecological competition, life histories, hybridization, and speciation), and consequently their systematics has been intensively investigated over several decades. Nevertheless, we lack a synthesis of relationships among the species. In the analyses reported here we use new DNA sequence data from the complete nuclear albumin gene (1818 bp) and the 12s mitochondrial gene (355 bp), as well as published data for four other genes (Wiens et al., 2006), up to a total of 6989 bp, to infer relationships. We relate these results to past systematic work based on morphology, allozymes, and DNA sequences. Although basal relationships show a strong consensus across studies, many terminal relationships remain in flux despite substantial sequencing and other molecular and morphological studies. This systematic instability appears to be a consequence of contemporaneous bursts of speciation in the late Miocene and Pliocene, yielding many closely related extant species in each of the four eastern species groups. Therefore we conclude that many relationships are likely to remain poorly resolved in the face of additional sequencing efforts. On the other hand, the current classification of the 45 eastern species into four species groups is supported. The Plethodon cinereus group (10 species) is the sister group to the clade comprising the other three groups, but these latter groups (Plethodon glutinosus [28 species], Plethodon welleri [5 species], and Plethodon wehrlei [2 species]) probably diverged from each other at approximately the same time. Copyright Â© 2012. Published by Elsevier Inc.
Mizuno, S; Andrews, C; Macgregor, H C
Intermediate repetitive sequences of Plethodon cinereus which comprised about 30% of the genomic DNA were isolated and iodinated with 125I. About 5% of the 125I-repetitive fraction hybridized with a large excess of DNA from P. dunni at Cot 20. About half of the 125I-DNA in the hybrids was resistant to extensive digestion with S-1 nuclease. The average molecular size of the S-1 nuclease-resistant fraction was about 100 nucleotide pairs. The melting temperature of the S-1 nuclease-resistant fraction was about 2 degrees lower than that of the corresponding fraction made with P. cinereus DNA. These results are taken to indicate the presence in the genomes of P. cinereus and P. dunni of evolutionarily stable "common" repetitive sequences. The average frequency of repetition of the common repetitive sequences is about 6,000 X in both species. The common repetitive fraction is also present in the genomes of other species of Plethodon, although the general populations of intermediate repetitive sequences are markedly different from one species to another. The cinereus--dunni common repetitive sequences could not be detected in plethodontids belonging to different tribes, nor in more distantly related amphibians. The profiles of binding of the common repetitive sequences to CsCl or CS2SO4-Ag+ density gradient fractions of P. dunni DNA suggested that these sequences consisted of heterogeneous components with respect to base compositions, and that they did not include large amounts of the genes for ribosomal RNA, 5S RNA, 4S RNA, or histone messenger RNA. In situ hybridization of the 3H-labelled intermediate repetitive sequences of P. cinereus to male meiotic chromosomes of the same species gave autoradiographs after an exposure of seven days showing all 14 chromosomes labelled. The pattern of labelling appeared not to be random, but was impossible to analyse on account of the irregular shapes and different degrees of stretching of diplotene and prometaphase chromosomes. In
Maerz, J C; Panebianco, N L; Madison, D M
Red-backed salamanders, Plethodon cinereus, show a variety of alarm responses to chemical cues from eastern garter snakes, Thamnophis sirtalis. We measured the foraging activity of red-backed salamanders exposed to water soiled by a garter snake (fed P. cinereus) or to unsoiled water. Salamanders exposed to snake-soiled water showed less foraging activity than salamanders exposed to unsoiled water; therefore, predators could have nonlethal effects on salamander populations. Our results also show additional factors influenced salamander foraging activity. Salamander foraging activity and responsiveness to chemical cues do not appear to have been affected by sex or food deprivation. Salamander foraging activity does appear to have been influenced by activity biorhythms. Foraging activity of animals in both treatments showed a bimodal periodicity that is consistent with natural activity patterns controlled by internal biorhythms. Exposure to snake-soiled water significantly reduced foraging activity during periods of peak foraging activity, but had a subtler effect on foraging activity during natural lulls in activity. We suggest that both activity biorhythms and exposure to chemical cues are important factors affecting salamander foraging behavior.
Ford, W. Mark; Mahoney, Kathleen R.; Russell, Kevin R.; Rodrigue, Jane L.; Riddle, Jason D.; Schuler, Thomas M.; Adams, Mary Beth
Forest management practices that mimic natural canopy disturbances, including prescribed fire and timber harvests, may reduce competition and facilitate establishment of favorable vegetative species within various ecosystems. Fire suppression in the central Appalachian region for almost a century has contributed to a transition from oak-dominated to more mesophytic, fire-intolerant forest communities. Prescribed fire coupled with timber removal is currently implemented to aid in oak regeneration and establishment but responses of woodland salamanders to this complex silvicultural system is poorly documented. The purpose of our research was to determine how woodland salamanders respond to shelterwood harvests following successive burns in a central Appalachian mixed-oak forest. Woodland salamanders were surveyed using coverboard arrays in May, July, and August–September 2011 and 2012. Surveys were conducted within fenced shelterwood-burn (prescribed fires, shelterwood harvest, and fencing to prevent white-tailed deer [Odocoileus virginianus] herbivory), shelterwood-burn (prescribed fires and shelterwood harvest), and control plots. Relative abundance was modeled in relation to habitat variables measured within treatments for mountain dusky salamanders (Desmognathus ochrophaeus), slimy salamanders (Plethodon glutinosus), and eastern red-backed salamanders (Plethodon cinereus). Mountain dusky salamander relative abundance was positively associated with canopy cover and there were significantly more individuals within controls than either shelterwood-burn or fenced shelterwood-burn treatments. Conversely, habitat variables associated with slimy salamanders and eastern red-backed salamanders did not differ among treatments. Salamander age-class structure within controls did not differ from shelterwood-burn or fenced shelterwood-burn treatments for any species. Overall, the woodland salamander assemblage remained relatively intact throughout the shelterwoodburn
MacNeil, Jami E; Williams, Rod N
Balancing timber production and conservation in forest management requires an understanding of how timber harvests affect wildlife species. Terrestrial salamanders are useful indicators of mature forest ecosystem health due to their importance to ecosystem processes and sensitivity to environmental change. However, the effects of timber harvests on salamanders, though often researched, are still not well understood. To further this understanding, we used artificial cover objects to monitor the relative abundance of terrestrial salamanders for two seasons (fall and spring) pre-harvest and five seasons post-harvest in six forest management treatments, and for three seasons post-harvest across the edge gradients of six recent clearcuts. In total, we recorded 19,048 encounters representing nine species of salamanders. We observed declines in mean encounters of eastern red-backed salamanders (Plethodon cinereus) and northern slimy salamanders (P. glutinosus) from pre- to post-harvest in group selection cuts and in clearcuts. However, we found no evidence of salamander declines at shelterwoods and forested sites adjacent to harvests. Edge effects induced by recent clearcuts influenced salamanders for approximately 20 m into the forest, but edge influence varied by slope orientation. Temperature, soil moisture, and canopy cover were all correlated with salamander counts. Our results suggest silvicultural techniques that remove the forest canopy negatively affect salamander relative abundance on the local scale during the years immediately following harvest, and that the depth of edge influence of clearcuts on terrestrial salamanders is relatively shallow (salamander populations across a forested landscape. Our results demonstrate the importance of examining species-specific responses and monitoring salamanders across multiple seasons and years. Long-term monitoring will be necessary to understand the full impacts of forest management on terrestrial salamanders.
Jami E MacNeil
Full Text Available Balancing timber production and conservation in forest management requires an understanding of how timber harvests affect wildlife species. Terrestrial salamanders are useful indicators of mature forest ecosystem health due to their importance to ecosystem processes and sensitivity to environmental change. However, the effects of timber harvests on salamanders, though often researched, are still not well understood. To further this understanding, we used artificial cover objects to monitor the relative abundance of terrestrial salamanders for two seasons (fall and spring pre-harvest and five seasons post-harvest in six forest management treatments, and for three seasons post-harvest across the edge gradients of six recent clearcuts. In total, we recorded 19,048 encounters representing nine species of salamanders. We observed declines in mean encounters of eastern red-backed salamanders (Plethodon cinereus and northern slimy salamanders (P. glutinosus from pre- to post-harvest in group selection cuts and in clearcuts. However, we found no evidence of salamander declines at shelterwoods and forested sites adjacent to harvests. Edge effects induced by recent clearcuts influenced salamanders for approximately 20 m into the forest, but edge influence varied by slope orientation. Temperature, soil moisture, and canopy cover were all correlated with salamander counts. Our results suggest silvicultural techniques that remove the forest canopy negatively affect salamander relative abundance on the local scale during the years immediately following harvest, and that the depth of edge influence of clearcuts on terrestrial salamanders is relatively shallow (<20 m. Small harvests (<4 ha and techniques that leave the forest canopy intact may be compatible with maintaining terrestrial salamander populations across a forested landscape. Our results demonstrate the importance of examining species-specific responses and monitoring salamanders across multiple
Dallalio, Eric A.; Brand, Adrianne; Grant, Evan H. Campbell
The distribution of the federally endangered Shenandoah Salamander (Plethodon shenandoah) is presumed to be limited by competition with the Red-backed Salamander (Plethodon cinereus). In particular, the current distribution of P. shenandoah is understood to be restricted to warmer and drier habitats because of interspecific interactions. These habitats may be particularly sensitive to climate change, though the influence of competition may also be affected by temperature and relative humidity. We investigated the response of P. shenandoah to competition with P. cinereus under four climate scenarios in 3-dimensional mesocosms. The results suggest that, although climate change may alleviate competitive pressure from P. cinereus, warmer temperatures may also significantly influence the persistence of the species across its known range.
Gillette; Jaeger; Peterson
Social monogamy, which does not necessarily imply mating or genetic monogamy, is important in the formation of male-female pair associations. We operationally define social monogamy as occurring when two heterosexual adults, exclusive of kin-directed behaviour, direct significantly less aggression and significantly more submission towards each other, and/or spend significantly more time associating with each other relative to other adult heterosexual conspecifics. Long-term pair associations (i.e. those lasting through a lengthy breeding season) that are characteristic of social monogamy are common in some taxa but are virtually unknown in amphibians. Recent studies, however, have suggested that red-backed salamanders, Plethodon cinereus, have complex (for amphibians) social systems. Our laboratory experiments tested the hypothesis that red-backed salamanders found in pairs in the forest display behaviours consistent with social monogamy. During the summer noncourtship season, newly collected male-females pairs showed no preference to associate with their partners more than with a novel conspecific of the opposite sex. However, during the autumn courtship season, paired males and females significantly directed preferential behaviours towards their partners rather than towards a surrogate or a novel paired salamander. Focal animals showed no significant preferences when presented with their partner and a novel single salamander, but they never directed preferential behaviours towards a novel salamander (whether paired or single) or a surrogate. These results are the first to suggest that a salamander species engages in social monogamy. Furthermore, our results suggest that social monogamy may not inhibit paired males and females from displaying alternative strategies: preferring partners when extrapair associations may be disadvantageous (i.e. the extrapair animal is already paired) but not preferring partners when extrapair associations may be advantageous (i
Placyk, John S; Graves, Brent M
While chemoreception is involved in a wide variety of salamander behaviors, the chemosensory system that mediates specific behaviors is rarely known. We investigated the role of the vomeronasal system (VNS) in foraging behavior of the red-backed salamander (Plethodon cinereus) by manipulating salamanders' abilities to detect nonvolatile chemical cues emitted by potential prey. Subjects received one of three treatments: (1) impaired vomeronasal system, (2) sham manipulation, and (3) no manipulation. The role of the VNS in mediating foraging on motile prey (Drosophila melanogaster) was investigated under three light conditions (bright, dim, dark). Salamanders with impaired VNSs foraged less efficiently than either of the other experimental groups by displaying the longest latency to attack and the lowest rate of prey capture, especially in the absence of visual cues. A second experiment utilized freshly killed prey to determine whether the VNS takes on added importance in the absence of visual or tactile cues associated with moving prey. Animals with impaired VNSs showed a decreased foraging efficiency on stationary prey under both dark and light conditions. In addition, a mark-recapture study of VNS-impaired and sham salamanders in the field also indicated that salamanders with impaired VNSs consumed fewer stationary prey compared to shams. The study indicates that the VNS plays a substantial role in the foraging behavior of the plethodontid salamander, P. cinereus.
Prey often avoid predator chemical cues, and in aquatic systems, prey may even appraise predation risk via cues associated with the predator's diet. However, this relationship has not been shown for terrestrial predator-prey systems, where the proximity of predators and prey, and the intensity of predator chemical cues in the environment, may be less than in aquatic systems. In the laboratory, we tested behavioural responses (avoidance, habituation and activity) of terrestrial red-backed salamanders, Plethodon cinereus, to chemical cues from garter snakes, Thamnophis sirtalis, fed either red-backed salamanders or earthworms (Lumbricus spp.). We placed salamanders in arenas lined with paper towels pretreated with snake chemicals, and monitored salamander movements during 120 min. Salamanders avoided substrates preconditioned by earthworm-fed (avoidanceX+/-SE=91.1+/-2.5%, N=25) and salamander-fed (95.2+/-2.5%, N=25) snakes, when tested against untreated substrate (control). Salamanders avoided cues from salamander-fed snakes more strongly (75.2+/-5.5%, N=25) than earthworm-fed snakes when subjected to both treatments simultaneously, implying that salamanders were sensitive to predator diet. Salamanders tended to avoid snake substrate more strongly during the last 60 min of a trial, but activity patterns were similar between salamanders exposed exclusively to control substrate versus those subject to snake cues. In another experiment, salamanders failed to avoid cues from dead conspecifics, suggesting that the stronger avoidance of salamander-fed snakes in the previous experiment was not directly due to chemical cues emitted by predator-killed salamanders. Salamanders also did not discriminate between cues from a salamander-fed snake versus a salamander-fed snake that was recently switched (i.e. <14 days) to an earthworm diet. Our results imply that terrestrial salamanders are sensitive to perceived predation risk via by-products of predator diet, and that snake
Fitzpatrick, Benjamin M; Allison, Amanda L
All animals and plants have intimate associations with microbes. Opinion has shifted from viewing microbes primarily as pathogens to the idea that healthy animals and plants carry specialized communities of coevolving microorganisms. However, the generality of this proposition is unknown because surveys rarely compare host-associated microbes with samples from relevant microhabitats. Symbiotic communities might be assembled from local environments with little evolutionary specialization. We evaluated the specificity of bacteria associated with salamander skin in comparison with surfaces in their immediate environments using 16S rRNA sequences. Host-associated and free-living samples were significantly different. However, relative abundances were strongly correlated; the most abundant taxa on salamander skin were also most abundant on moist debris on the forest floor. Thus, although bacterial assemblages on salamander skin are statistically differentiated from those on inanimate surfaces, they are not entirely 'distinct'. Candidate salamander specialists were few in number and occurred at low relative abundances. Within some OTUs, differences in allele frequency suggested genetic specialization at finer levels. Although host-associated and free-living assemblages were similar, a range of more or less specialized symbiotes was evident and bacteria on salamander skin were often specific genotypes of OTUs commonly found on other moist surfaces in the environment.
Margolis, Ellis; Malevich, Steven B.
Anthropogenic alteration of ecosystem processes confounds forest management and conservation of rare, declining species. Restoration of forest structure and fire hazard reduction are central goals of forest management policy in the western United States, but restoration priorities and treatments have become increasingly contentious. Numerous studies have documented changes in fire regimes, forest stand structure and species composition following a century of fire exclusion in dry, frequent-fire forests of the western U.S. (e.g., ponderosa pine and dry mixed-conifer). In contrast, wet mixed-conifer forests are thought to have historically burned infrequently with mixed- or high-severity fire—resulting in reduced impacts from fire exclusion and low restoration need—but data are limited. In this study we quantified the current forest habitat of the federally endangered, terrestrial Jemez Mountains salamander (Plethodon neomexicanus) and compared it to dendroecological reconstructions of historical habitat (e.g., stand structure and composition), and fire regime parameters along a gradient from upper ponderosa pine to wet mixed-conifer forests. We found that current fire-free intervals in Jemez Mountains salamander habitat (116–165 years) are significantly longer than historical intervals, even in wet mixed-conifer forests. Historical mean fire intervals ranged from 10 to 42 years along the forest gradient. Low-severity fires were historically dominant across all forest types (92 of 102 fires). Although some mixed- or highseverity fire historically occurred at 67% of the plots over the last four centuries, complete mortality within 1.0 ha plots was rare, and asynchronous within and among sites. Climate was an important driver of temporal variability in fire severity, such that mixed- and high-severity fires were associated with more extreme drought than low-severity fires. Tree density in dry conifer forests historically ranged from open (90 trees/ha) to
Hess, Alexandra; McAllister, Caroline; DeMarchi, Joseph; Zidek, Makenzie; Murone, Julie; Venesky, Matthew D
Immune function is a costly line of defense against parasitism. When infected with a parasite, hosts frequently lose mass due to these costs. However, some infected hosts (e.g. highly resistant individuals) can clear infections with seemingly little fitness losses, but few studies have tested how resistant hosts mitigate these costly immune defenses. We explored this topic using eastern red-backed salamanders Plethodon cinereus and the fungal pathogen Batrachochytrium dendrobatidis (Bd). Bd is generally lethal for amphibians, and stereotypical symptoms of infection include loss in mass and deficits in feeding. However, individuals of P. cinereus can clear their Bd infections with seemingly few fitness costs. We conducted an experiment in which we repeatedly observed the feeding activity of Bd-infected and non-infected salamanders. We found that Bd-infected salamanders generally increased their feeding activity compared to non-infected salamanders. The fact that we did not observe any differences in mass change between the treatments suggests that increased feeding might help Bd-infected salamanders minimize the costs of an effective immune response.
Anthony, Carl D; Venesky, Matthew D; Hickerson, Cari-Ann M
1. When studying speciation, researchers commonly examine reproductive isolation in recently diverged populations. Polymorphic species provide an opportunity to examine the role of reproductive isolation in populations that may be in the process of divergence. 2. We examined a polymorphic population of Plethodon cinereus (red-backed salamanders) for evidence of sympatric ecological separation by colour morphology. Recent studies have correlated temperature and climate with colour morphology in this species, but no studies have looked at differences in diet or mate choice between colour morphs. We used artificial cover objects to assess salamander diet, mating preference and surface activity over a 2-year period at a field site in north-eastern Ohio. 3. We detected differences in diet between two colour morphs, striped and unstriped. The diets of striped individuals were significantly more diverse and were made up of more profitable prey than the diets of unstriped salamanders. 4. Opposite sex pairs were made up of individuals of the same colour morph and striped males were found more often with larger females than were unstriped males. 5. We corroborate findings of earlier studies suggesting that the unstriped form is adapted to warmer conditions. Unstriped individuals were the first to withdraw from the forest floor as temperatures fell in the late fall. We found no evidence that the colour morphs responded differently to abiotic factors such as soil moisture and relative humidity, and responses to surface temperatures were also equivocal. 6. We conclude that the two colour morphs exhibit some degree of ecological separation and tend to mate assortatively, but are unlikely to be undergoing divergence given the observed frequency of intermorph pairings.
Grant, Alexa H; Liebgold, Eric B
Behavioral traits can be influenced by predation rates of color morphs, potentially leading to reduced boldness or increased escape behaviors in one color morph. The red-backed salamander, Plethodon cinereus, is a small terrestrial salamander whose color morphs have different diets and select different microhabitats, but little is known about potential differences in dispersal behaviors. We used fine-scale genetic spatial autocorrelation to examine 122 P. cinereus in a color-polymorphic population at 10 microsatellite loci in order to generate estimates of spatial genetic structure for each color morph. Differences in spatial genetic structure have been used extensively to infer within-population sex-biased dispersal but have never been used to test for dispersal differences between other groups within populations such as color morphs. We found evidence for color-biased dispersal, but not sex-biased dispersal. Striped salamanders had significant positive genetic structure in the shortest distance classes indicating philopatry. In contrast, unstriped salamanders showed a lack of spatial genetic structure at shorter distances and higher than expected genetic similarity at further distances, as expected if they are dispersing from their natal site. These results show that genetic methods typically used for sex-biased dispersal can be used to investigate differences in dispersal between morphs that vary discretely in polymorphic populations, such as color morphs. © The American Genetic Association 2017. All rights reserved. For permissions, please e-mail: firstname.lastname@example.org.
US Fish and Wildlife Service, Department of the Interior — On July 8, 2009 a night area search was conducted for presence of Cheat Mountain salamanders (Plethodon nettingi) on the Kelly Elkins Tract of the Refuge. A total of...
Damien B Wilburn
Full Text Available In response to pervasive sexual selection, protein sex pheromones often display rapid mutation and accelerated evolution of corresponding gene sequences. For proteins, the general dogma is that structure is maintained even as sequence or function may rapidly change. This phenomenon is well exemplified by the three-finger protein (TFP superfamily: a diverse class of vertebrate proteins co-opted for many biological functions - such as components of snake venoms, regulators of the complement system, and coordinators of amphibian limb regeneration. All of the >200 structurally characterized TFPs adopt the namesake "three-finger" topology. In male red-legged salamanders, the TFP pheromone Plethodontid Modulating Factor (PMF is a hypervariable protein such that, through extensive gene duplication and pervasive sexual selection, individual male salamanders express more than 30 unique isoforms. However, it remained unclear how this accelerated evolution affected the protein structure of PMF. Using LC/MS-MS and multidimensional NMR, we report the 3D structure of the most abundant PMF isoform, PMF-G. The high resolution structural ensemble revealed a highly modified TFP structure, including a unique disulfide bonding pattern and loss of secondary structure, that define a novel protein topology with greater backbone flexibility in the third peptide finger. Sequence comparison, models of molecular evolution, and homology modeling together support that this flexible third finger is the most rapidly evolving segment of PMF. Combined with PMF sequence hypervariability, this structural flexibility may enhance the plasticity of PMF as a chemical signal by permitting potentially thousands of structural conformers. We propose that the flexible third finger plays a critical role in PMF:receptor interactions. As female receptors co-evolve, this flexibility may allow PMF to still bind its receptor(s without the immediate need for complementary mutations. Consequently
Wilburn, Damien B; Bowen, Kathleen E; Doty, Kari A; Arumugam, Sengodagounder; Lane, Andrew N; Feldhoff, Pamela W; Feldhoff, Richard C
In response to pervasive sexual selection, protein sex pheromones often display rapid mutation and accelerated evolution of corresponding gene sequences. For proteins, the general dogma is that structure is maintained even as sequence or function may rapidly change. This phenomenon is well exemplified by the three-finger protein (TFP) superfamily: a diverse class of vertebrate proteins co-opted for many biological functions - such as components of snake venoms, regulators of the complement system, and coordinators of amphibian limb regeneration. All of the >200 structurally characterized TFPs adopt the namesake "three-finger" topology. In male red-legged salamanders, the TFP pheromone Plethodontid Modulating Factor (PMF) is a hypervariable protein such that, through extensive gene duplication and pervasive sexual selection, individual male salamanders express more than 30 unique isoforms. However, it remained unclear how this accelerated evolution affected the protein structure of PMF. Using LC/MS-MS and multidimensional NMR, we report the 3D structure of the most abundant PMF isoform, PMF-G. The high resolution structural ensemble revealed a highly modified TFP structure, including a unique disulfide bonding pattern and loss of secondary structure, that define a novel protein topology with greater backbone flexibility in the third peptide finger. Sequence comparison, models of molecular evolution, and homology modeling together support that this flexible third finger is the most rapidly evolving segment of PMF. Combined with PMF sequence hypervariability, this structural flexibility may enhance the plasticity of PMF as a chemical signal by permitting potentially thousands of structural conformers. We propose that the flexible third finger plays a critical role in PMF:receptor interactions. As female receptors co-evolve, this flexibility may allow PMF to still bind its receptor(s) without the immediate need for complementary mutations. Consequently, this unique
Cosentino, Bradley J; Moore, Jean-David; Karraker, Nancy E; Ouellet, Martin; Gibbs, James P
Evolutionary change has been demonstrated to occur rapidly in human-modified systems, yet understanding how multiple components of global change interact to affect adaptive evolution remains a critical knowledge gap. Climate change is predicted to impose directional selection on traits to reduce thermal stress, but the strength of directional selection may be mediated by changes in the thermal environment driven by land use. We examined how regional climatic conditions and land use interact to affect genetically based color polymorphism in the eastern red-backed salamander (Plethodon cinereus). P. cinereus is a woodland salamander with two primary discrete color morphs (striped, unstriped) that have been associated with macroclimatic conditions. Striped individuals are most common in colder regions, but morph frequencies can be variable within climate zones. We used path analysis to analyze morph frequencies among 238,591 individual salamanders across 1,170 sites in North America. Frequency of striped individuals was positively related to forest cover in populations occurring in warmer regions (>7°C annually), a relationship that was weak to nonexistent in populations located in colder regions (≤7°C annually). Our results suggest that directional selection imposed by climate warming at a regional scale may be amplified by forest loss and suppressed by forest persistence, with a mediating effect of land use that varies geographically. Our work highlights how the complex interaction of selection pressures imposed by different components of global change may lead to divergent evolutionary trajectories among populations.
Jessica A. Wooten; William B. Sutton; Thomas K. Pauley
Increased availability of habitat and climate data has facilitated much research concerning the influence of these characteristics on the structure of salamander communities. We aimed to outline environmental requirements influencing the distribution of three sympatric plethodontid salamander species, including Plethodon cinereus, Desmognathus ochrophaeus...
Liebgold, Eric B; Cabe, Paul R; Jaeger, Robert G; Leberg, Paul L
In the majority of birds and mammals, social monogamy is not congruent with genetic monogamy. No research to date has compared social and genetic monogamy in amphibians. We analysed paternity in clutches of red-backed salamanders (Plethodon cinereus), a species in which social monogamy has been demonstrated in the laboratory, and 28% of individuals in the forest are found in male-female pairs in the noncourtship season. We collected 16 clutches of eggs of P. cinereus in the southern Appalachian Mountains of Virginia and collected tail clippings from attending mothers. We genotyped embryos and adults at five microsatellite loci in order to analyse paternity of clutches. Most clutches (84.6%) had multiple sires, with two to three sires per clutch. In this study, 25% of clutches had males in addition to females attending eggs. None of the mothers of these clutches were genetically monogamous. All attending males sired some of the offspring in the clutch that they attended (between 9% and 50%) but never sired a majority in that clutch. We conclude that, at least in this population, social monogamy in P. cinereus is not concomitant with genetic monogamy.
Lisa Ollivier; Hartwell H. Welsh Jr
Life stage determination for many western plethodontids often requires dissection of the specimen. Availability of reliable external measures that could be applied under field conditions would enhance future studies of the genus Plethodon. We examined preserved specimens of the Del Norte Salamander, Plethodon elongatus, taken from...
Walker, Donald M; Lawrence, Brandy R; Esterline, Dakota; Graham, Sean P; Edelbrock, Michael A; Wooten, Jessica A
The flow of energy within an ecosystem can be considered either top-down, where predators influence consumers, or bottom-up, where producers influence consumers. Plethodon cinereus (Red-backed Salamander) is a terrestrial keystone predator who feeds on invertebrates within the ecosystem. We investigated the impact of the removal of P. cinereus on the detritivore food web in an upland deciduous forest in northwest Ohio, U.S.A. A total of eight aluminum enclosures, each containing a single P. cinereus under a small log, were constructed in the deciduous forest. On Day 1 of the experiment, four salamanders were evicted from four of the eight enclosures. Organic matter and soil were collected from the center of each enclosure at Day 1 and Day 21. From each sample, DNA was extracted, fungal-specific amplification performed, and 454 pyrosequencing was used to sequence the nuclear ribosomal internal transcribed spacer (ITS2) region and partial ribosomal large subunit (LSU). Changes in overall fungal community composition or species diversity were not statistically significant between treatments. Statistically significant shifts in the most abundant taxonomic groups of fungi were documented in presence but not absence enclosures. We concluded that P. cinereus does not affect the overall composition or diversity of fungal communities, but does have an impact on specific groups of fungi. This study used a metagenomics-based approach to investigate a missing link among a keystone predator, P. cinereus, invertebrates, and fungal communities, all of which are critical in the detritivore food web.
Richard M. DeGraaf; Mariko. Yamasaki
Terrestrial salamanders are sensitive to forest disturbance associated with even-aged management. We studied the distribution of redback salamanders (Plethodon cinereus) for 4 yr at edges between even-aged northern hardwood stands along three replicate transects in each of three edge contrast types: regeneration/mature, sapling/mature, and...
Richard M. DeGraaf; Mariko. Yamasaki
Salamanders are abundant vertebrates in many forest ecosystems, and their annual biomass production can be important in forest food webs (Pough et al. 1987). Population densities of eastern redback salamanders (Plethodon cinereus) can exceed 2 individuals/m2 in deciduous forests of the United States (Heatwole 1962, Jaeger 1980...
US Fish and Wildlife Service, Department of the Interior — Cheat Mountain Salamanders (Plethodon nettingi) are known to occur in only 5 counties and the western edge of Grant County along the Allegheny Front in the eastern...
Newman, Catherine E; Gregory, T Ryan; Austin, Christopher C
The genus Plethodon is the most species-rich salamander genus in North America, and nearly half of its species face an uncertain future. It is also one of the most diverse families in terms of genome sizes, which range from 1C = 18.2 to 69.3 pg, or 5-20 times larger than the human genome. Large genome size in salamanders results in part from accumulation of transposable elements and is associated with various developmental and physiological traits. However, genome sizes have been reported for only 25% of the species of Plethodon (14 of 55). We collected genome size data for Plethodon serratus to supplement an ongoing phylogeographic study, reconstructed the evolutionary history of genome size in Plethodontidae, and inferred probable genome sizes for the 41 species missing empirical data. Results revealed multiple genome size changes in Plethodon: genomes of western Plethodon increased, whereas genomes of eastern Plethodon decreased, followed by additional decreases or subsequent increases. The estimated genome size of P. serratus was 21 pg. New understanding of variation in genome size evolution, along with genome size inferences for previously unstudied taxa, provide a foundation for future studies on the biology of plethodontid salamanders.
William B. Sutton
Full Text Available Global climate change represents one of the most extensive and pervasive threats to wildlife populations. Amphibians, specifically salamanders, are particularly susceptible to the effects of changing climates due to their restrictive physiological requirements and low vagility; however, little is known about which landscapes and species are vulnerable to climate change. Our study objectives included, (1 evaluating species-specific predictions (based on 2050 climate projections and vulnerabilities to climate change and (2 using collective species responses to identify areas of climate refugia for conservation priority salamanders in the northeastern United States. All evaluated salamander species were projected to lose a portion of their climatic niche. Averaged projected losses ranged from 3%–100% for individual species, with the Cow Knob Salamander (Plethodon punctatus, Cheat Mountain Salamander (Plethodon nettingi, Shenandoah Mountain Salamander (Plethodon virginia, Mabee’s Salamander (Ambystoma mabeei, and Streamside Salamander (Ambystoma barbouri predicted to lose at least 97% of their landscape-scale climatic niche. The Western Allegheny Plateau was predicted to lose the greatest salamander climate refugia richness (i.e., number of species with a climatically-suitable niche in a landscape patch, whereas the Central Appalachians provided refugia for the greatest number of species during current and projected climate scenarios. Our results can be used to identify species and landscapes that are likely to be further affected by climate change and potentially resilient habitats that will provide consistent climatic conditions in the face of environmental change.
Chinnadurai, Sathya K; Cooper, David; Dombrowski, Daniel S; Poore, Matthew F; Levy, Michael G
Chytridiomycosis is an often fatal fungal disease of amphibians caused by Batrachochytrium dendrobatidis. This disease has been implicated in the worldwide decline of many anuran species, but studies of chytridiomycosis in wild salamanders are limited. Between August 2006 and December 2006, we tested wild amphibians in North Carolina, USA (n=212) by polymerase chain reaction (PCR). We identified three PCR-positive animals: one Rana clamitans and two Plethodontid salamanders. We experimentally infected two species of native North Carolina Plethodontid salamanders, the slimy salamander (Plethodon glutinosus) and the Blue Ridge Mountain dusky salamander (Desmognathus orestes) with 1,000,000 zoospores of B. dendrobatidis per animal. Susceptibility was species dependent; all slimy salamanders developed clinical signs of chytridiomycosis, and one died, whereas dusky salamanders remained unaffected. In a second experiment, we challenged naïve slimy salamanders with either 10,000 or 100,000 motile zoospores per animal. Clinical signs consistent with chytridiomycosis were not observed at either dose or in uninfected controls during the 45 days of this experiment. All animals inoculated with B. dendrobatidis in both experiments, regardless of dose, tested positive by PCR. Our study indicates that slimy salamanders are more susceptible to clinical chytridiomycosis than dusky salamanders, and in a laboratory setting, a dose greater than 100,000 zoospores per animal is required to induce clinical disease. This study also indicates that PCR is a very sensitive tool for detecting B. dendrobatidis infection, even in animals that are clinically unaffected, thus positive results should be interpreted with caution.
Muletz Wolz, Carly R; Yarwood, Stephanie A; Campbell Grant, Evan H; Fleischer, Robert C; Lips, Karen R
The amphibian skin microbiome is recognized for its role in defence against pathogens, including the deadly fungal pathogen Batrachochytrium dendrobatidis (Bd). Yet, we have little understanding of evolutionary and ecological processes that structure these communities, especially for salamanders and closely related species. We investigated patterns in the distribution of bacterial communities on Plethodon salamander skin across host species and environments. Quantifying salamander skin microbiome structure contributes to our understanding of how host-associated bacteria are distributed across the landscape, among host species, and their putative relationship with disease. We characterized skin microbiome structure (alpha-diversity, beta-diversity and bacterial operational taxonomic unit [OTU] abundances) using 16S rRNA gene sequencing for co-occurring Plethodon salamander species (35 Plethodon cinereus, 17 Plethodon glutinosus, 10 Plethodon cylindraceus) at three localities to differentiate the effects of host species from environmental factors on the microbiome. We sampled the microbiome of P. cinereus along an elevational gradient (n = 50, 700-1,000 m a.s.l.) at one locality to determine whether elevation predicts microbiome structure. Finally, we quantified prevalence and abundance of putatively anti-Bd bacteria to determine if Bd-inhibitory bacteria are dominant microbiome members. Co-occurring salamanders had similar microbiome structure, but among sites salamanders had dissimilar microbiome structure for beta-diversity and abundance of 28 bacterial OTUs. We found that alpha-diversity increased with elevation, beta-diversity and the abundance of 17 bacterial OTUs changed with elevation (16 OTUs decreasing, 1 OTU increasing). We detected 11 putatively anti-Bd bacterial OTUs that were present on 90% of salamanders and made up an average relative abundance of 83% (SD ± 8.5) per salamander. All salamanders tested negative for Bd. We conclude that
Assessment of potentially asymmetrical characters (such as fighting ability and resident advantage) is often important in determining the outcome of agonistic interactions. Loss of body parts, a predator defence mechanism used by many animals, may lead to a reduction in fighting ability and may be easily assessed by competitors. We investigated the influence of tail loss on the expression of agonistic behaviour in the territorial red-backed salamander, Plethodon cinereus. Residents and intruders were matched for body size, and pairs were tested in all combinations of tailed or tailless residents with tailed or tailless intruders. Neither residents nor intruders altered their behaviour based on their own tail condition, but they did alter their behaviour based on the tail condition of their opponents. Intruders showed more aggression or less submission towards tailless residents than towards tailed residents. When contests were between residents and intruders of the same tail condition (both tailed or tailless), intruders were more aggressive towards residents when both were tailless than when both were tailed, indicating that tail loss does not directly hamper aggressive displays. In contests where the asymmetry between residents and intruders was small (based on tail condition and residency status), intruders showed more aggression and less submission than in contests where the asymmetry was large. Residents did not differ in their behaviour for most comparisons. Thus, for intruders, the tail condition of residents is an important determinant of agonistic behaviour displayed in territorial contests. For residents, factors other than tail condition (such as resource value) may be more likely to influence their behaviour. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.
Webster, T. Preston
In some populations of the salamander Plethodon cinereus, two polymorphic esterase loci are in linkage disequilibrium. Short-term stability of the linkage disequilibrium is demonstrated by an age class analysis. Long, perhaps very long, term stability is suggested by its distribution. This stability and concordant geographic variation in allelic frequencies imply selective origin and maintenance. Data on the frequencies of two color morphs suggest that formation of the linkage disequilibrium is dependent on the genetic background. Images PMID:4515614
Burton, T.M.; Likens, G.E.
There were about 2950 salamanders per ha (1770 g/ha wet wt) in the Hubbard Brook Experimental Forest in New Hampshire. The terrestrial species, Plethodon cinereus, accounted for about 93.5% of the total biomass while the streamside species, Desmognathus fuscus, Eurycea bislineata and Gyrinophilus porphyriticus, accounted for the remaining 6.5%. Notophthalmus viridescens was present, but was rare and insignificant in the biomass calculations. The population size of salamanders at Hubbard Brook appears to be stable. The biomass of salamanders is about twice that of birds during the bird's peak (breeding) season and is about equal to the biomass of small mammals.
Byrne, G C; Pemberton, P J
A case of red-back spider (Latrodectus mactans hasselti) envenomation in a neonate is presented. The signs of envenomation in a neonate are discussed, together with the dose of antivenom necessary for the successful treatment of envenomation. In episodes of untractable crying in neonates, the possibility that the baby has been bitten by a red-back spider should be considered.
Ilseman, Kelly; Hoffmann, Kristine
On a spring morning in Maine, traps made of nets rise above vernal pools in a small wetland, ready to collect salamanders. The traps were designed by groups of rural and urban high school students from Maine and Massachusetts participating in the University of Maine Upward Bound Math Science Program (UBMS) at the university campus in Orono, Maine.…
Adams, D C; Rohlf, F J
Ecological character displacement describes a pattern where morphological differences between sympatric species are enhanced through interspecific competition. Although widely considered a pervasive force in evolutionary ecology, few clear-cut examples have been documented. Here we report a case of ecological character displacement between two salamander species, Plethodon cinereus and Plethodon hoffmani. Morphology was quantified by using linear measurements and landmark-based geometric morphometric methods for specimens from allopatric and sympatric populations from two geographic transects in south-central Pennsylvania, and stomach contents were assayed to quantify food resource use. Morphological variation was also assessed in 13 additional allopatric populations. In both transects, we found significant morphological differentiation between sympatric populations that was associated with a reduction in prey consumption in sympatry and a segregation of prey according to prey size. No trophic morphological or resource use differences were found between allopatric populations, and comparisons of sympatric populations with randomly paired allopatric populations revealed that the observed sympatric morphological differentiation was greater than expected by chance. The major trophic anatomical differences between sympatric populations relates to functional and biomechanical differences in jaw closure: sympatric P. hoffmani have a faster closing jaw, whereas sympatric P. cinereus have a slower, stronger jaw. Because salamanders immobilize prey of different sizes in different ways, and because the observed sympatric biomechanical differences in jaw closure are associated with the differences in prey consumption, the observed character displacement has a functional ecological correlate, and we can link changes in form with changes in function in this apparent example of character displacement.
Salamander * Plethodon cinereus Spring Peeper * Hyla crucifer American Toad Bufo americanus Gray Treefrog * Hyla versicolor Green Frog Rana clamitans Wood...Red Back Salamander ( Plethodon cinereus ) and the Wood Frog (Rana sylvatica) were also observed utilizing this tidal freshwater habitat. The high salt...K68 Table K46. Mammals Observed in the Study Area Wetlands Common Name Scientific Name Opossum Didelphis marsupia.is Masked Shrew * Sorex cinereus
Ransom, Tami S
Ecosystem engineers create habitat that can be used by other species in multiple ways, such as refugees from predators, places to breed, or areas with increased prey resources. I conducted a series of enclosure experiments to: (1) determine if salamanders use earthworm burrows, and (2) examine the potential influence of earthworm burrow use and indirect effects on salamander intra- and interspecific competition, predator avoidance, and seasonal performance. I found that one species of woodland salamander, Plethodon cinereus, used earthworm burrows 50% of the time when burrows were present. Neither adults nor juveniles of the congeneric P. glutinosus used earthworm burrows. Intraspecific, but not interspecific, competition by P. cinereus affected salamander behavior when earthworms were absent, with P. cinereus found under cover objects >70% of the time when alone or with a P. glutinosus, but only 40% of the time when with another P. cinereus. When earthworms were present, the behavior of P. cinereus was similar across salamander treatments. Earthworms decreased the amount of leaf litter and microinvertebrates, although this did not affect salamander mass. In subsequent experiments using only P. cinereus, the refuge provided by earthworm burrows increased the survival of P. cinereus over the winter and allowed P. cinereus to avoid being consumed by the common garter snake (Thamnophis sirtalis). Because earthworm burrows provide a refuge for P. cinereus during intraspecific encounters, in the presence of a predator and over the winter, they may serve as an important belowground-aboveground linkage in eastern forests where salamanders are common.
Lemaître, Jérôme; Fortin, Daniel; Montiglio, Pierre-Olivier; Darveau, Marcel
Parasites can play an important role in the dynamics of host populations, but empirical evidence remains sparse. We investigated the role of bot fly (Cuterebra spp.) parasitism in red-backed voles (Myodes gapperi) by first assessing the impacts of the parasite on the probability of vole survival under stressful conditions as well as on the reproductive activity of females. We then identified the main factors driving both the individual risk of infection and the abundance of bot flies inside red-backed voles. Finally, we evaluated the impacts of bot fly prevalence on the growth rate of vole populations between mid-July and mid-August. Thirty-six populations of red-backed voles were sampled in the boreal forest of Québec, Canada. The presence and the abundance of parasites in voles, two host life history traits (sex and body condition), three indices of habitat complexity (tree basal area, sapling basal area, coarse woody debris volume), and vole abundance were considered in models evaluating the effects of bot flies on host populations. We found that the probability of survival of red-backed voles in live traps decreased with bot fly infection. Both the individual risk of infection and the abundance of bot flies in red-backed voles were driven mainly by vole abundance rather than by the two host life history traits or the three variables of habitat complexity. Parasitism had population consequences: bot fly prevalence was linked to a decrease in short-term growth rate of vole populations over the summer. We found that bot flies have the potential to reduce survival of red-backed voles, an effect that may apply to large portions of populations.
Muletz-Wolz, Carly R.; DiRenzo, Graziella V.; Yarwood, Stephanie A.; Grant, Evan H. Campbell; Fleischer, Robert C.; Lips, Karen R.
Diverse bacteria inhabit amphibian skin; some of those bacteria inhibit growth of the fungal pathogen Batrachochytrium dendrobatidis. Yet there has been no systematic survey of anti-B. dendrobatidis bacteria across localities, species, and elevations. This is important given geographic and taxonomic variations in amphibian susceptibility to B. dendrobatidis. Our collection sites were at locations within the Appalachian Mountains where previous sampling had indicated low B. dendrobatidis prevalence. We determined the numbers and identities of anti-B. dendrobatidis bacteria on 61 Plethodon salamanders (37 P. cinereus, 15 P. glutinosus, 9 P. cylindraceus) via culturing methods and 16S rRNA gene sequencing. We sampled co-occurring species at three localities and sampled P. cinereus along an elevational gradient (700 to 1,000 meters above sea level [masl]) at one locality. We identified 50 anti-B. dendrobatidis bacterial operational taxonomic units (OTUs) and found that the degree of B. dendrobatidis inhibition was not correlated with relatedness. Five anti-B. dendrobatidis bacterial strains occurred on multiple amphibian species at multiple localities, but none were shared among all species and localities. The prevalence of anti-B. dendrobatidis bacteria was higher at Shenandoah National Park (NP), VA, with 96% (25/26) of salamanders hosting at least one anti-B. dendrobatidis bacterial species compared to 50% (7/14) at Catoctin Mountain Park (MP), MD, and 38% (8/21) at Mt. Rogers National Recreation Area (NRA), VA. At the individual level, salamanders at Shenandoah NP had more anti-B. dendrobatidis bacteria per individual (μ = 3.3) than those at Catoctin MP (μ = 0.8) and at Mt. Rogers NRA (μ = 0.4). All salamanders tested negative for B. dendrobatidis. Anti-B. dendrobatidis bacterial species are diverse in central Appalachian Plethodon salamanders, and their distribution varied geographically. The antifungal bacterial species that we identified may play a protective
Feder, M E
In preliminary experiments, salamanders of three species (Desmognathus ochrophaeus, Plethodon cinereus and Plethodon jordani) required 5-11 days to complete metabolic acclimation to a constant warm temperature; the rate of oxygen consumption (VO2) decreased 16-28% during acclimation. Unfed animals of each species underwent cyclic exposure to 5 and 21 degrees C at three different cycle periods (12 hr, 4-5 days, 51 days), or constant exposure to 14 degrees C for 102 days. The experimental treatments significantly affected the VO2 measured at 5, 14, 17.5 and 21 degrees C. The direction and magnitude of the acclimatory effects upon VO2 were inconsistent among species and among experimental temperatures, and resulted in little energy saving. The VO2 during exposure to cyclic temperatures averaged only 83% of that during preliminary experiments, perhaps as a response to starvation.
Svendsen, Jannie K.; Sell, Henrik; Bøcher, Peder Klith;
In this study we evaluate nest site and habitat preferences of Red-backed Shrike (Lanius collurio) in the national park Mols Bjerge, Denmark. In total, 28 nests were found during 2010 and 2011. To determine nesting site preferences, the species composition of vegetation in scrub used for nesting...... was compared to that of nearest unused scrub. To evaluate habitat preferences, a Resource Selection Probability Function (RSPF) was modelled based on presence/absence data. The habitat factors were represented by Light Detection And Ranging (LIDAR) derived measures of vegetation height and topographic wetness...... wetness, and that it was unaffected by the distance to the nearest road/path. These results provide some guidelines for management, showing that Red-backed Shrike for nesting preferred habitats with high wetness, possibly linked to food availability, and heterogeneous vegetation, consistent...
Tøttrup, Anders P.; Pedersen, Lykke; Onrubia, Alejandro;
e current Northern Hemisphere migration systems are believed to have arisen since the last glaciation. In many cases, birds do not migrate strait from breeding to non-breeding areas but ﬂy via a detour. All western European populations of red-backed shrikes Lanius collurio are assumed to reach...... their southern African wintering grounds detouring via southeast Europe. Based on theoretical considerations under an optimality framework this detour is apparently optimal. Here, we use individual geolocator data on red-backed shrikes breeding in Spain to show that these birds do indeed detour via southeast...... migration the situation was quite diﬀerent with the detour apparently being optimal. However, when considering potential wind assistance estimated total air distances during autumn migration were overall similar and the barrier crossing shorter along the observed routes. We conclude that considering...
Kraemer, Andrew C; Adams, Dean C
In Batesian mimicry a palatable mimic deceives predators by resembling an unpalatable model. The evolution of Batesian mimicry relies on the visual capabilities of the potential predators, as prey detection provides the selective force driving evolutionary change. We compared the visual capabilities of several potential predators to test predictions stemming from the hypothesis of Batesian mimicry between two salamanders: the model species Notophthalmus viridescens, and polymorphic mimic, Plethodon cinereus. First, we found mimicry to be restricted to coloration, but not brightness. Second, only bird predators appeared able to discriminate between the colors of models and nonmimic P. cinereus. Third, estimates of salamander conspicuousness were background dependent, corresponding to predictions only for backgrounds against which salamanders are most active. These results support the hypothesis that birds influence the evolution of Batesian mimicry in P. cinereus, as they are the only group examined capable of differentiating N. viridescens and nonmimetic P. cinereus. Additionally, patterns of conspicuousness suggest that selection from predators may drive the evolution of conspicuousness in this system. This study confirms the expectation that the visual abilities of predators may influence the evolution of Batesian mimicry, but the role of conspicuousness may be more complex than previously thought. © 2013 The Author(s). Evolution © 2013 The Society for the Study of Evolution.
Jung, R.E.; Droege, S.; Sauer, J.R.; Landy, R.B.
In response to concerns about amphibian declines, a study evaluating and validating amphibian monitoring techniques was initiated in Shenandoah and Big Bend National Parks in the spring of 1998. We evaluate precision, bias, and efficiency of several sampling methods for terrestrial and streamside salamanders in Shenandoah National Park and assess salamander abundance in relation to environmental variables, notably soil and water pH. Terrestrial salamanders, primarily redback salamanders (Plethodon cinereus), were sampled by searching under cover objects during the day in square plots (10 to 35 m2). We compared population indices (mean daily and total counts) with adjusted population estimates from capture-recapture. Analyses suggested that the proportion of salamanders detected (p) during sampling varied among plots, necessitating the use of adjusted population estimates. However, adjusted population estimates were less precise than population indices, and may not be efficient in relating salamander populations to environmental variables. In future sampling, strategic use of capture-recapture to verify consistency of p's among sites may be a reasonable compromise between the possibility of bias in estimation of population size and deficiencies due to inefficiency associated with the estimation of p. The streamside two-lined salamander (Eurycea bislineata) was surveyed using four methods: leaf litter refugia bags, 1 m2 quadrats, 50 x 1 m visual encounter transects, and electric shocking. Comparison of survey methods at nine streams revealed congruent patterns of abundance among sites, suggesting that relative bias among the methods is similar, and that choice of survey method should be based on precision and logistical efficiency. Redback and two-lined salamander abundance were not significantly related to soil or water pH, respectively.
Full Text Available Roads are a pervasive feature in the landscape, and their ecological effects on vertebrate wildlife have been well documented. The main types of effect described are mainly negative consequences on birds and other vertebrates. The major impact of roads on birds includes habitat fragmentation, traffic noise and direct mortality from road kills. However, some passerines, such as the Red-backed Shrike, seem to often use areas close to roads for nesting and hunting purposes.The aim of this paper is to study the importance of road proximity for the selection of suitable shrubs for nesting by the Red-backed Shrike in the farmland landscapes of central Italy. To achieve this goal, the hierarchical partitioning procedures of Generalized Linear Models (GLM are applied in order to quantify the relative effects of a number of independent variables.At least 77% of the nests we identified were positioned less than 25 meters from roads. The mean distance from the nest to the nearest road was 12.9 ± 18.1 m. The analysis of the relative importance of each variable revealed that “road distance” is one of variables most associated with nesting probability in suitable shrubs. It is reasonable to argue that suitable shrubs and the presence of open spaces for hunting, both of which exist close to countryside roads, might represent the favourable components of the breeding habitat selections of Red-backed Shrikes. Our results can provide useful indications for census techniques and for the planning of conservation measures for the species in agricultural landscapes.
Hocking, Daniel; Babbitt, Kimberly; Yamasaki, Mariko
Poster presented at the 98th annual meeting of the Ecological Society of America (ESA) in Minneapolis, Minnesota, USA. We used a non-linear, parametric model accounting for detection probability to quantify red-backed salamander (Plethodon cinereus) abundance across clearcut-forest edges. This approach allows for projection across landscapes and prediction given alternative logging plans.
Connette, Grant M; Semlitsch, Raymond D
Forest management often represents a balance between social, economic, and ecological objectives. In the eastern United States, numerous studies have established that terrestrial salamander populations initially decline in abundance following timber harvest, yet the large-scale and long-term consequences are relatively unknown. We used count data from terrestrial survey points to examine the relation between salamander abundance and historic timber harvest while accounting for imperfect detection of individuals. Overall, stream- and terrestrial-breeding salamanders appeared to differ by magnitude of population decline, rate of population recovery, and extent of recolonization from surrounding forest. Specifically, estimated abundance of both species groups was positively associated with stand age and recovery rates were predicted to increase over time for red-legged salamanders (Plethodon shermani) and decrease in stream-breeding species. Abundance of stream-breeding salamanders was predicted to reach a peak by 100 years after timber harvest, and the population growth rate of red-legged salamanders was predicted to undergo a significant increase 100 years after harvest. Estimated abundance of stream-breeding salamanders in young forest stands was also negatively associated with the distance to adjacent forest, a result that suggests immigration has a role in the recovery of these species. Our results indicate that salamander abundance in young forest stands may be only modestly lower than in more mature forest but that full recovery from timber harvest may take a substantial amount of time and that species life history may affect patterns of recovery. Historia de Vida como un Vaticinador de la Tasa de Recuperación de una Salamandra a la Colecta de Madera en los Bosques del Sur de los Apalaches, E.U.A.
Matthew H Becker
Full Text Available Chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis (Bd, is an infectious disease that causes population declines of many amphibians. Cutaneous bacteria isolated from redback salamanders, Plethodon cinereus, and mountain yellow-legged frogs, Rana muscosa, inhibit the growth of Bd in vitro. In this study, the bacterial community present on the skin of P. cinereus individuals was investigated to determine if it provides protection to salamanders from the lethal and sub-lethal effects of chytridiomycosis. When the cutaneous bacterial community was reduced prior to Bd exposure, salamanders experienced a significantly greater decrease in body mass, which is a symptom of the disease, when compared to infected individuals with a normal bacterial community. In addition, a greater proportion of infected individuals with a reduced bacterial community experienced limb-lifting, a behavior seen only in infected individuals. Overall, these results demonstrate that the cutaneous bacterial community of P. cinereus provides protection to the salamander from Bd and that alteration of this community can change disease resistance. Therefore, symbiotic microbes associated with this species appear to be an important component of its innate skin defenses.
Adam L. Crane
Full Text Available When new predators invade a habitat, either through range extensions or introductions, prey may be at a high risk because they do not recognize the predators as dangerous. The nine-banded armadillo (Dasypus novemcinctus has recently expanded its range in North America. Armadillos forage by searching soil and leaf litter, consuming invertebrates and small vertebrates, including salamanders. We tested whether Ozark zigzag salamanders (Plethodon angusticlavius from a population coexisting with armadillos for about 30 years exhibit antipredator behavior in the presence of armadillo chemical cues and whether they can discriminate between stimuli from armadillos and a nonpredatory sympatric mammal (white-tailed deer, Odocoileus virginianus. Salamanders appeared to recognize substrate cues from armadillos as a threat because they increased escape behaviors and oxygen consumption. When exposed to airborne cues from armadillos, salamanders also exhibited an antipredator response by spending more time in an inconspicuous posture. Additionally, individually consistent behaviors across treatments for some response variables suggest the potential for a behavioral syndrome in this species.
Becker, Matthew H; Harris, Reid N
Chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis (Bd), is an infectious disease that causes population declines of many amphibians. Cutaneous bacteria isolated from redback salamanders, Plethodon cinereus, and mountain yellow-legged frogs, Rana muscosa, inhibit the growth of Bd in vitro. In this study, the bacterial community present on the skin of P. cinereus individuals was investigated to determine if it provides protection to salamanders from the lethal and sub-lethal effects of chytridiomycosis. When the cutaneous bacterial community was reduced prior to Bd exposure, salamanders experienced a significantly greater decrease in body mass, which is a symptom of the disease, when compared to infected individuals with a normal bacterial community. In addition, a greater proportion of infected individuals with a reduced bacterial community experienced limb-lifting, a behavior seen only in infected individuals. Overall, these results demonstrate that the cutaneous bacterial community of P. cinereus provides protection to the salamander from Bd and that alteration of this community can change disease resistance. Therefore, symbiotic microbes associated with this species appear to be an important component of its innate skin defenses.
Deban, Stephen M; Scales, Jeffrey A
Low temperature reduces the performance of muscle-powered movements, but in movements powered by elastic recoil mechanisms, this effect can be mitigated and performance can be increased. To better understand the morphological basis of high performance and thermal robustness of elastically powered movements, we compared feeding dynamics at a range of temperatures (5-25°C) in two species of terrestrial plethodontid salamanders, Plethodon metcalfi and Ensatina eschscholtzii, which differ in tongue muscle architecture and the mechanism of tongue projection. We found that Ensatina is capable of ballistic projection with a mean muscle mass-specific power of 2100 W kg(-1), revealing an elastic mechanism. Plethodon, in contrast, projected its tongue non-ballistically with a mean power of only 18 W kg(-1), indicating it is muscle powered. Ensatina projected its tongue significantly farther than Plethodon and with dynamics that had significantly lower thermal sensitivity at temperatures below 15°C. These performance differences were correlated with morphological differences, namely elongated collagenous aponeuroses in the projector muscle of Ensatina as compared with Plethodon, which are likely the site of energy storage, and the absence in Ensatina of projector muscle fibers attaching to the tongue skeleton that allows projection to be truly ballistic. These findings demonstrate that, in these otherwise similar species, the presence in one species of elaborated connective tissue in series with myofibers confers not only 10-fold greater absolute performance but also greater thermal robustness of performance. We conclude that changes in muscle and connective tissue architecture are sufficient to alter significantly the mechanics, performance and thermal robustness of musculoskeletal systems.
Lauer, Antje; Simon, Mary Alice; Banning, Jenifer L; Lam, Brianna A; Harris, Reid N
Among the microbiota of amphibian skin are bacteria that produce antifungal compounds. We isolated cutaneous bacteria from the skins of three populations of the nest-attending plethodontid salamander Hemidactylium scutatum and subsequently tested the bacterial isolates against two different fungi (related to Mariannaea elegans and Rhizomucor variabilis) that were obtained from dead salamander eggs. The culturable antifungal bacteria were phylogenetically characterized based on 16S rRNA phylogeny, and belonged to four phyla, comprising 14 bacterial families, 16 genera and 48 species. We found that about half of the antifungal bacterial genera and families were shared with a related salamander species, but there was virtually no overlap at the species level. The proportion of culturable antifungal bacterial taxa shared between two large populations of H. scutatum was the same as the proportion of taxa shared between H. scutatum and Plethodon cinereus, suggesting that populations within a species have unique antifungal bacterial species. Approximately 30% of individuals from both salamander species carried anti-M. elegans cutaneous bacteria and almost 90% of P. cinereus and 100% of H. scutatum salamanders carried anti-R. variabilis cutaneous bacteria. A culture independent method (PCR/DGGE) revealed a shared resident bacterial community of about 25% of the entire resident bacterial community within and among populations of H. scutatum. Thus, the culturable antifungal microbiota was far more variable on salamander skins than was the bacterial microbiota detected by PCR/DGGE. The resident cutaneous antifungal bacteria may play an important role in amphibians' innate defense against pathogens, including the lethal chytrid fungus Batrachochytrium dendrobatidis.
Svendsen, Jannie K.; Sell, Henrik; Bøcher, Peder Klith;
In this study we evaluate nest site and habitat preferences of Red-backed Shrike (Lanius collurio) in the national park Mols Bjerge, Denmark. In total, 28 nests were found during 2010 and 2011. To determine nesting site preferences, the species composition of vegetation in scrub used for nesting...... was compared to that of nearest unused scrub. To evaluate habitat preferences, a Resource Selection Probability Function (RSPF) was modelled based on presence/absence data. The habitat factors were represented by Light Detection And Ranging (LIDAR) derived measures of vegetation height and topographic wetness...... as well as distance to nearest road/path, as an indicator of human disturbance. Scrub used as nesting sites were characterized by thorny shrub species such as Prunus spinosa and Rubus fruticosus. RSPF showed that shrike presence was positively correlated with vegetation heterogeneity and high topographic...
Muletz-Wolz, Carly R; DiRenzo, Graziella V; Yarwood, Stephanie A; Campbell Grant, Evan H; Fleischer, Robert C; Lips, Karen R
Diverse bacteria inhabit amphibian skin; some of those bacteria inhibit growth of the fungal pathogen Batrachochytrium dendrobatidis Yet there has been no systematic survey of anti-B. dendrobatidis bacteria across localities, species, and elevations. This is important given geographic and taxonomic variations in amphibian susceptibility to B. dendrobatidis Our collection sites were at locations within the Appalachian Mountains where previous sampling had indicated low B. dendrobatidis prevalence. We determined the numbers and identities of anti-B. dendrobatidis bacteria on 61 Plethodon salamanders (37 P. cinereus, 15 P. glutinosus, 9 P. cylindraceus) via culturing methods and 16S rRNA gene sequencing. We sampled co-occurring species at three localities and sampled P. cinereus along an elevational gradient (700 to 1,000 meters above sea level [masl]) at one locality. We identified 50 anti-B. dendrobatidis bacterial operational taxonomic units (OTUs) and found that the degree of B. dendrobatidis inhibition was not correlated with relatedness. Five anti-B. dendrobatidis bacterial strains occurred on multiple amphibian species at multiple localities, but none were shared among all species and localities. The prevalence of anti-B. dendrobatidis bacteria was higher at Shenandoah National Park (NP), VA, with 96% (25/26) of salamanders hosting at least one anti-B. dendrobatidis bacterial species compared to 50% (7/14) at Catoctin Mountain Park (MP), MD, and 38% (8/21) at Mt. Rogers National Recreation Area (NRA), VA. At the individual level, salamanders at Shenandoah NP had more anti-B. dendrobatidis bacteria per individual (μ = 3.3) than those at Catoctin MP (μ = 0.8) and at Mt. Rogers NRA (μ = 0.4). All salamanders tested negative for B. dendrobatidis Anti-B. dendrobatidis bacterial species are diverse in central Appalachian Plethodon salamanders, and their distribution varied geographically. The antifungal bacterial species that we identified may play a protective
Sessions, Stanley K
Salamanders (Amphibia: Caudata/Urodela) have been the subject of numerous cytogenetic studies, and data on karyotypes and genome sizes are available for most groups. Salamanders show a more-or-less distinct dichotomy between families with large chromosome numbers and interspecific variation in chromosome number, relative size, and shape (i.e. position of the centromere), and those that exhibit very little variation in these karyological features. This dichotomy is the basis of a major model of karyotype evolution in salamanders involving a kind of 'karyotypic orthoselection'. Salamanders are also characterized by extremely large genomes (in terms of absolute mass of nuclear DNA) and extensive variation in genome size (and overall size of the chromosomes), which transcends variation in chromosome number and shape. The biological significance and evolution of chromosome number and shape within the karyotype is not yet understood, but genome size variation has been found to have strong phenotypic, biogeographic, and phylogenetic correlates that reveal information about the biological significance of this cytogenetic variable. Urodeles also present the advantage of only 10 families and less than 600 species, which facilitates the analysis of patterns within the entire order. The purpose of this review is to present a summary of what is currently known about overall patterns of variation in karyology and genome size in salamanders. These patterns are discussed within an evolutionary context.
I examined the potential influence of climate change on the dynamics of a previously studied hybrid zone between a pair of terrestrial salamanders at the Coweeta Hydrologic Laboratory, U.S. Forest Service, in the Nantahala Mountains of North Carolina, USA. A 16-year study led by Nelson G. Hairston, Sr. revealed that Plethodon teyahalee and Plethodon shermani hybridized at intermediate elevations, forming a cline between 'pure' parental P. teyahalee at lower elevations and 'pure' parental P. shermani at higher elevations. From 1974 to 1990 the proportion of salamanders at the higher elevation scored as 'pure' P. shermani declined significantly, indicating that the hybrid zone was spreading upward. To date there have been no rigorous tests of hypotheses for the movement of this hybrid zone. Using temperature and precipitation data from Coweeta, I re-analyzed Hairston's data to examine whether the observed elevational shift was correlated with variation in either air temperature or precipitation from the same time period. For temperature, my analysis tracked the results of the original study: the proportion of 'pure' P. shermani at the higher elevation declined significantly with increasing mean annual temperature, whereas the proportion of 'pure' P. teyahalee at lower elevations did not. There was no discernable relationship between proportions of 'pure' individuals of either species with variation in precipitation. From 1974 to 1990, low-elevation air temperatures at the Coweeta Laboratory ranged from annual means of 11.8 to 14.2 °C, compared with a 55-year average (1936-1990) of 12.6 °C. My re-analyses indicate that the upward spread of the hybrid zone is correlated with increasing air temperatures, but not precipitation, and provide an empirical test of a hypothesis for one factor that may have influenced this movement. My results aid in understanding the potential impact that climate change may have on the ecology and evolution of terrestrial salamanders in
Hemerik, L.; Geertsma, M.; Waasdorp, S.; Middelveld, R.P.; Kleef, van H.; Klok, T.C.
Populations of many bird species strongly declined in Western Europe in the late twentieth century. One such species is the Red-backed Shrike in the Netherlands. In one of the last strongholds of this species, the Bargerveen Reserve, the breeding population flourished in the 1990s due to rewetting m
Tianlong FU; Shan SUN; Chaonan ZHU; Yanfu QU
The red-backed ratsnake (Elaphe rufodorsata) is widely distributed in East Asia, especially China. This species is a common snake in plain river network region.In the past several decades,E. rufodorsata has dramatically declined due to the effect of human activities and over hunting for traditional Chinese medicine. We developed nine species-specific microsatellite loci in 190 individuals collected from Huzhou, Zhejiang province in China. These markers revealed a high degree of genetic diversity (13–41 alleles per locus) and heterozygosity (HO ranged from 0.266 to 0.941, and HE ranged from 0.851 to 0.937). No locus exhibited significant deviations from Hardy–Weinberg equilibrium. There was no evidence of linkage disequilibrium among pairs of loci. These microsatellite markers were described in our study will be valuable tools for the long term management and population-level studies (e.g. the population structure, genetic diversity and variation, individual paternity and evolutionary history) of the species.
Brand, Adrianne; Wiewel, Amber N. M.; Grant, Evan H. Campbell
Natural gas production from the Marcellus shale is rapidly increasing in the northeastern United States. Most of the endemic terrestrial salamander species in the region are classified as ‘globally secure’ by the IUCN, primarily because much of their ranges include state- and federally protected lands, which have been presumed to be free from habitat loss. However, the proposed and ongoing development of the Marcellus gas resources may result in significant range restrictions for these and other terrestrial forest salamanders. To begin to address the gaps in our knowledge of the direct impacts of shale gas development, we developed occurrence models for five species of terrestrial plethodontid salamanders found largely within the Marcellus shale play. We predicted future Marcellus shale development under several scenarios. Under scenarios of 10,000, 20,000, and 50,000 new gas wells, we predict 4%, 8%, and 20% forest loss, respectively, within the play. Predictions of habitat loss vary among species, but in general, Plethodon electromorphus and Plethodonwehrlei are predicted to lose the greatest proportion of forested habitat within their ranges if future Marcellus development is based on characteristics of the shale play. If development is based on current well locations,Plethodonrichmondi is predicted to lose the greatest proportion of habitat. Models showed high uncertainty in species’ ranges and emphasize the need for distribution data collected by widespread and repeated, randomized surveys.
Doty, Kari A; Wilburn, Damien B; Bowen, Kathleen E; Feldhoff, Pamela W; Feldhoff, Richard C
Gene co-option is a major force in the evolution of novel biological functions. In plethodontid salamanders, males deliver proteinaceous courtship pheromones to the female olfactory system or transdermally to the bloodstream. Molecular studies identified three families of highly duplicated, rapidly evolving pheromones (PRF, PMF, and SPF). Analyses for Plethodon salamanders revealed pheromone mixtures of primarily PRF and PMF. The current study demonstrates that in Desmognathus ocoee--a plesiomorphic species with transdermal delivery--SPF is the major pheromone component representing >30% of total protein. Chromatographic profiles of D. ocoee pheromones were consistent from May through October. LC/MS-MS analysis suggested uniform SPF isoform expression between individual male D. ocoee. A gene ancestry for SPF with the Three-Finger Protein superfamily was supported by intron-exon boundaries, but not by the disulfide bonding pattern. Further analysis of the pheromone mixture revealed paralogs to peptide hormones that contained mutations in receptor binding regions, such that these novel molecules may alter female physiology by acting as hormone agonists/antagonists. Cumulatively, gene co-option, duplication, and neofunctionalization have permitted recruitment of additional gene families for pheromone activity. Such independent co-option events may be playing a key role in salamander speciation by altering male traits that influence reproductive success. Copyright © 2015 Elsevier B.V. All rights reserved.
Graudins, Andis; Little, Michelle J; Pineda, Sandy S; Hains, Peter G; King, Glenn F; Broady, Kevin W; Nicholson, Graham M
The venom of the European black widow spider Latrodectus tredecimguttatus (Theridiidae) contains several high molecular mass (110-140 kDa) neurotoxins that induce neurotransmitter exocytosis. These include a vertebrate-specific α-latrotoxin (α-LTX-Lt1a) responsible for the clinical symptoms of latrodectism and numerous insect-specific latroinsectoxins (LITs). In contrast, little is known about the expression of these toxins in other Latrodectus species despite the fact that envenomation by these spiders induces a similar clinical syndrome. Here we report highly conserved α-LTX, α-LIT and δ-LIT sequence tags in Latrodectus mactans, Latrodectus hesperus and Latrodectus hasselti venoms using tandem mass spectrometry, following bioassay-guided separation of venoms by liquid chromatography. Despite this sequence similarity, we show that the anti-α-LTX monoclonal antibody 4C4.1, raised against α-LTX-Lt1a, fails to neutralize the neurotoxicity of all other Latrodectus venoms tested in an isolated chick biventer cervicis nerve-muscle bioassay. This suggests that there are important structural differences between α-LTXs in theridiid spider venoms. We therefore cloned and sequenced the α-LTX from the Australian red-back spider L. hasselti (α-LTX-Lh1a). The deduced amino acid sequence of the mature α-LTX-Lh1a comprises 1180 residues (∼132kDa) with ∼93% sequence identity with α-LTX-Lt1a. α-LTX-Lh1a is composed of an N-terminal domain and a central region containing 22 ankyrin-like repeats. The presence of two furin cleavage sites, conserved with α-LTX-Lt1a, indicates that α-LTX-Lh1a is derived from the proteolytic cleavage of an N-terminal signal peptide and C-terminal propeptide region. However, we show that α-LTX-Lh1a has key substitutions in the 4C4.1 epitope that explains the lack of binding of the monoclonal antibody.
Full Text Available Background: Due to importance and fatal affect of Red-back spiders, Latrodectus hasselti, a faunistic survey for presence of this spider in Bandar Abbas has been conducted. This animal is considerably the most medically importance spiders all over the world.Methods: Live adult spider specimens were collected from Bandar Abbas town using hand catch conventional method and transferred to the laboratory throughout the summer of 2008. They were identified based on morphological characteristics and taxonomic keys and confirmed by some external experts.Results: Results showed the occurrence of the red-back spider, L. hasselti from Bandar Abbas, southern port of Iran. Two female specimens were found. The spider had specific morphological characters including black color with an obvious orange to red longitudinal strip on its upper parts of abdomen. Conclusion: Although the specimens were collected from south of the country, however since the region is an important harbor and port and goods come form different parts of world we assume the possibility of arrival from its origin and native breeding sites of the world. Therefore further investigation is needed to clarify the presence of this species in different parts of Iran.
Townsend, Jason M; Driscoll, Charles T; Rimmer, Christopher C; McFarland, Kent P
High-elevation ecosystems of the northeastern United States are vulnerable to deposition and environmental accumulation of atmospheric pollutants, yet little work has been done to assess mercury (Hg) concentrations in organisms occupying montane ecosystems. The authors present data on Hg concentrations in ground-foraging insectivorous songbirds, a terrestrial salamander, and forest floor horizons sampled along a forested elevational gradient from 185 m to 1273 m in the Catskill Mountains, New York, USA. Mean Hg concentrations in Catharus thrushes and the salamander Plethodon cinereus increased with elevation, as did Hg concentrations in all forest floor horizons. Mean Hg concentrations in organic soils at approximately 1200 m elevation (503.5 ± 17.7 ng/g, dry wt) were 4.4-fold greater than those at approximately 200 m. Montane ecosystems of the northeastern United States, and probably elsewhere, are exposed to higher levels of atmospheric Hg deposition as reflected in accumulation patterns in the forest floor and associated high-elevation fauna. This information can be used to parameterize and test Hg transport and bioaccumulation models of landscape-specific patterns and may serve as a monitoring tool for decision makers considering future controls on Hg emissions. Further investigation is needed into the potential effects of increased Hg concentrations on high-elevation fauna. © 2013 SETAC.
Heuring, Whitney L; Mathis, Alicia
Learning to use a landmark as a beacon to locate resources is one of the simplest forms of spatial learning. We tested whether landmark learning occurs in a semifossorial salamander that migrates annually to breeding ponds as adults. Juvenile spotted salamanders (Ambystoma maculatum) were tested in square containers with a plastic feeding dish in each corner, and a piece of earthworm was placed in one randomly-chosen dish. For landmark-trained salamanders, a rock was placed beside the dish containing the prey. For control salamanders, the rock was placed beside a randomly selected feeding dish. Each salamander was trained once every 2 days for 30 days. Significantly more landmark-trained salamanders than control salamanders entered the landmark area first, and landmark-trained individuals had faster latencies to enter the landmark area and longer stay-times. These results suggest that spotted salamanders are able to locate resources by associating their positions with landmarks.
Pedersen, Lykke; Geertsma, Marten; Tøttrup, Anders P.
When breeding, food availability is essential for optimal reproductive output and is potentially one of the main factors limiting breeding success, especially in single brooded long-distance migratory birds. In this study, we examined the diet (as a measure of prey availability) of two Red...... on a larger variety of species in colder summers. Adults had a more diverse diet and generally fed on smaller prey items than did young birds. Thus, age- and environment-related differences must be taken into account when describing the diet of theRed-backed Shrike.Direct nest observations produced different...... results for diet composition than did nest and pellet samples, underlining the importance of using different methods in diet assessments. Detailed knowledge on limiting factors on the breeding grounds, such as food availability, is crucial for mitigating population declines of vulnerable species...
Jaeger, R G
Plethodon richmondi shenandoah occurs in at least three geographically isolated talus slopes in Shenandoah National Park, Virginia, U.S.A., each surrounded by a continuous population of Plethodon c. cinereus in the soil outside the talus. Distributions are contiguous but largely non-overlapping. The talus presents a much drier habitat than does the surrounding soil. Four experiments were designed to test the responses of the two species to moisture and substrate. Although shenandoah lives in a habitat generally drier than that of cinereus, both species choose the wet end of a moisture gradient and do not differ significantly in moisture preference. When given choices between a substrate of rock or soil, the two species respond similarly: neither expresses a preference when both substrates are moist and both choose soil over rock as the substrates dry, showing that substrate preference is based on moisture content and not texture. A third experiment demonstrates that cinereus suffers significantly greater mortality and loss of body water when subjected to a drying rock substrate than when subjected to a soil substrate, since the latter holds moisture longer. Thus the talus most likely presents a greater stress of dehydration to salamanders than does the soil. A fourth experiment shows that when forced to dehydrate, shenandoah survives longer, loses significantly less body water per hour, and withstands a greater loss of body water before death than does cinereus.The conclusions drawn are that cinereus inhabits areas of deep soil not due to a preference for that substrate but due to the requirement of a moist substrate, and it cannot enter the talus due to the dry conditions there. P. r. shenandoah, on the other hand, neither prefers the rocky nor the dry conditions of the talus and is probably excluded from the soil by the presence of cinereus. The survival of shenandoah in the talus is due, at least in part, to its ability to withstand the stress of dehydration for
Jefferson, Dale M; Ferrari, Maud CO; Mathis, Alicia; Hobson, Keith A.; Eric R Britzke; Crane, Adam L; Blaustein, Andrew R.; Chivers, Douglas P.
Introduction Many species of ambystomatid salamanders are dependent upon highly variable temporary wetlands for larval development. High larval densities may prompt the expression of a distinct head morphology that may facilitate cannibalism. However, few studies have characterized structural cannibalism within natural populations of larval salamanders. In this study we used two species of larval salamanders, long-toed (Ambystoma macrodactylum) and ringed salamanders (A. annulatum). Head morp...
Němec, Michal; Syrová, Michaela; Dokoupilová, Lenka; Veselý, Petr; Šmilauer, Petr; Landová, Eva; Lišková, Silvie; Fuchs, Roman
We compared the responses of the nesting red-backed shrikes (Lanius collurio) to three dummies of a common nest predator, the Eurasian jay (Garrulus glandarius), each made from a different material (stuffed, plush, and silicone). The shrikes performed defensive behaviour including attacks on all three dummies. Nevertheless, the number of attacks significantly decreased from the stuffed dummy through the plush dummy and finally to the silicone dummy. Our results show that wild birds use not only colours but also other surface features as important cues for recognition and categorization of other bird species. Moreover, the silicone dummy was attacked only when presented after the stuffed or plush dummy. Thus, we concluded that the shrikes recognized the jay only the stuffed (with feathered surface) and plush (with hairy surface) dummies during the first encounter. Recognition of the silicon dummy (with glossy surface) was facilitated by previous encounters with the more accurate model. This process resembles the effect of perceptual priming, which is widely described in the literature on humans.
Atakuziev, Bakhadir U; Wright, Christine E; Graudins, Andis; Nicholson, Graham M; Winkel, Kenneth D
We describe the first Steatoda capensis envenomation treated with CSL red-back spider antivenom (RBSAV). The patient, a 51-year-old female, developed acute local pain, swelling, redness, and diaphoresis in association with tender lymphadenopathy and hypertension. These features responded, in a dose-dependent manner, to RBSAV. In vitro studies confirmed that RBSAV could neutralize S. capensis venom at equivalent concentrations required to neutralize red-back spider (Latrodectus hasselti) venom. Similar data were obtained using Mexican Latrodectus mactans antivenom (Aracmyn®). Although S. capensis yielded similar quantities of venom protein as L. hasselti, pooled S. capensis and Steatoda grossa venom was more rapidly toxic to insects than either L. hasselti or Latrodectus tredecimguttatus venom. By contrast, both Latrodectus venoms were more potent than S. capensis venom in contracting rat isolated mesenteric arteries. Size-exclusion and anion-exchange chromatography was used to purify a 130 kDa fraction from S. capensis venom that induced contracture and loss of twitch tension in chick isolated biventer cervicis nerve-muscle preparations in a manner similar to α-latrotoxin. This activity was abolished by pre-incubation with RBSAV. We conclude that 'steatodism' may overlap more closely with latrodectism than previously recognized and that this bite should be managed in the same way as for Australian red-back envenomation.
... purposes of collecting biological data Jemez Mountain salamanders (Plethodon neomexicanus) within New...; construction monitoring; capture and relocation of Jemez Mountain salamanders ] (Plethodon neomexicanus) within... temporarily hold for the purposes of collecting biological data Jemez Mountain salamanders...
Gao, Ke-Qin; Shubin, Neil H
Salamanders are a model system for studying the rates and patterns of the evolution of new anatomical structures. Recent discoveries of abundant Late Jurassic and Early Cretaceous salamanders are helping to address these issues. Here we report the discovery of well-preserved Middle Jurassic salamanders from China, which constitutes the earliest known record of crown-group urodeles (living salamanders and their closest relatives). The new specimens are from the volcanic deposits of the Jiulongshan Formation (Bathonian), Inner Mongolia, China, and represent basal members of the Cryptobranchidae, a family that includes the endangered Asian giant salamander (Andrias) and the North American hellbender (Cryptobranchus). These fossils document a Mesozoic record of the Cryptobranchidae, predating the previous record of the group by some 100 million years. This discovery provides evidence to support the hypothesis that the divergence of the Cryptobranchidae from the Hynobiidae had taken place in Asia before the Middle Jurassic period.
Fisher-Reid, M Caitlin; Engstrom, Tag N; Kuczynski, Caitlin A; Stephens, Patrick R; Wiens, John J
Speciation is often categorized based on geographic modes (allopatric, parapatric or sympatric). Although it is widely accepted that species can arise in allopatry and then later become sympatrically or parapatrically distributed, patterns in the opposite direction are also theoretically possible (e.g. sympatric lineages or ecotypes becoming parapatric), but such patterns have not been shown at a macrogeographic scale. Here, we analyse genetic, climatic, ecological and morphological data and show that two typically sympatric colour morphs of the salamander Plethodon cinereus (redback and leadback) appear to have become parapatrically distributed on Long Island, New York, with pure-redback populations in the west and pure-leadback populations in the east (and polymorphic populations in between and on the mainland). In addition, the pure-leadback populations in eastern Long Island are genetically, ecologically and morphologically divergent from both mainland and other Long Island populations, suggesting the possibility of incipient speciation. This parapatric separation seems to be related to the different ecological preferences of the two morphs, preferences which are present on the mainland and across Long Island. These results potentially support the idea that spatial segregation of sympatric ecotypes may sometimes play an important part in parapatric speciation. © 2013 John Wiley & Sons Ltd.
Harris, Reid N; Lauer, Antje; Simon, Mary Alice; Banning, Jenifer L; Alford, Ross A
Chytridiomycosis, caused by the skin fungus Batrachochytrium dendrobatidis (Bd), has caused population declines of many amphibians in remote protected habitats. Progress has been made in understanding the pathogen's life cycle, documenting its devastating effects on individual amphibians and on populations, and understanding how and why disease outbreaks occur. No research has directly addressed the critical question of how to prevent declines and extinctions caused by outbreaks of the disease. We have identified a number of bacterial species of amphibian skin that inhibit Bd in vitro. Here, we demonstrate that a species of anti-Bd skin bacteria can be successfully added to skins of salamanders Plethodon cinereus, and that addition of this bacterium reduced the severity of a disease symptom in experimentally infected individuals. This is the first demonstration that manipulating the natural skin microbiota of an amphibian species can alter the pathogen's negative effects on infected amphibians and appears to be the first demonstration that an epibiotic manipulation of any wildlife species can lessen the effects of an emerging infectious disease. It suggests that probiotic or bio-augmentation manipulations of cutaneous microbiota could have the potential to reduce susceptibility of amphibians to the disease in nature. This is the first approach suggested that could slow or halt epidemic outbreaks and allow successful reintroductions of amphibian species that have become locally or globally extinct in the wild. Our results also suggest a mechanism for the association of climate change and the likelihood of chytridiomycosis outbreaks via the effects of the former on antifungal bacterial communities.
Kraemer, A C; Serb, J M; Adams, D C
Conspicuousness, or having high contrast relative to the surrounding background, is a common feature of unpalatable species. Several hypotheses have been proposed to explain the occurrence of conspicuousness, and while most involve the role of conspicuousness as a direct signal of unpalatability to potential predators, one hypothesis suggests that exaggerated conspicuousness may evolve in unpalatable species to reduce predator confusion with palatable species (potential Batesian mimics). This hypothesis of antagonistic coevolution between palatable and unpalatable species hinges on the 'cost of conspicuousness', in which conspicuousness increases the likelihood of predation more in palatable species than in unpalatable species. Under this mimicry scenario, four patterns are expected: (i) mimics will more closely resemble local models than models from other localities, (ii) there will be a positive relationship between mimic and model conspicuousness, (iii) models will be more conspicuous in the presence of mimics, and (iv) when models and mimics differ in conspicuousness, mimics will be less conspicuous than models. We tested these predictions in the salamander mimicry system involving Notophthalmus viridescens (model) and one colour morph of Plethodon cinereus (mimic). All predictions were supported, indicating that selection for Batesian mimicry not only influences the evolution of mimics, but also the evolution of the models they resemble. These findings indicate that mimicry plays a large role in the evolution of model warning signals, particularly influencing the evolution of conspicuousness. © 2015 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2015 European Society For Evolutionary Biology.
Glorioso, Brad M.; Waddle, Hardin; Hefner, Jeromi
The Spotted Salamander is a wide-ranging salamander of the eastern United States that typically breeds in winter or early spring in ephemeral pools in lowland forests. Ambystoma maculatum is known to deposit 2-4 egg masses per year, each containing 1-250 eggs. As part of ongoing research into the ecology and reproductive biology of Spotted Salamanders in the Kisatchie District of Kisatchie National Forest in Natchitoches Parish, Louisiana, USA, we have been counting the number of embryos per egg mass. We captured seven female A. maculatum in a small pool, six of which were still gravid. We took standard measurements, including SVL, and then implanted a Passive Integrated Transponder (PIT tag) into each adult female as was the protocol. About an hour after processing these animals we marked new A. maculatum egg masses found in the same small pool using PVC pin flags pushed carefully through the outer jelly. We did not have enough time to process them that evening, and it was not until a few days later that we photographed those masses. We discovered that one of the masses contained a PIT tag in the outer jelly that corresponded to one of the six gravid females that were marked that same evening. To our knowledge, this is the first report of PIT tags being the means, albeit coincidentally, by which a particular egg mass of Ambystoma maculatum has been assigned to a particular female. For our purposes, losing the PIT tag from the adult female is counter to the goals of our study of this population, and we will no longer be implanting PIT tags into gravid females.
Jefferson, Dale M; Ferrari, Maud Co; Mathis, Alicia; Hobson, Keith A; Britzke, Eric R; Crane, Adam L; Blaustein, Andrew R; Chivers, Douglas P
Many species of ambystomatid salamanders are dependent upon highly variable temporary wetlands for larval development. High larval densities may prompt the expression of a distinct head morphology that may facilitate cannibalism. However, few studies have characterized structural cannibalism within natural populations of larval salamanders. In this study we used two species of larval salamanders, long-toed (Ambystoma macrodactylum) and ringed salamanders (A. annulatum). Head morphometrics and stable isotopic values of carbon (δ(13)C) and nitrogen (δ(15)N) were used to identify the presence or absence of structural cannibalism. Weather conditions were also analyzed as a potential factor associated with the expression of cannibalistic morphology. Populations of salamander larvae did not consistently exhibit cannibalistic morphologies throughout collection periods. Larval long-toed salamanders exhibited trophic polymorphisms when relatively lower precipitation amounts were observed. Larval ringed salamanders were observed to be cannibalistic but did not exhibit polymorphisms in this study. Structural cannibalism may be transient in both species; however in long-toed salamanders this morphology is necessary for cannibalism. Ringed salamanders can be cannibalistic without morphological adaptations; however the cannibal morph may prolong the viable time period for cannibalism. Additionally, weather conditions may alter pond hydroperiod, subsequently influencing head morphology and cannibalism.
Vinikour, W. S.; LaGory, K. E.; Adduci, J. J.; Environmental Science Division
The purpose of this conservation assessment is to summarize existing knowledge regarding the biology and ecology of the Siskiyou Mountains salamander and Scott Bar salamander, identify threats to the two species, and identify conservation considerations to aid federal management for persistence of the species. The conservation assessment will serve as the basis for a conservation strategy for the species.
Regulatory agencies need appropriate indicators of stream permanence to aid in jurisdictional determinations for headwater streams. We evaluated salamanders as permanence indicators because they are often abundant in fishless headwaters. Salamander and habitat data were collect...
Sun, Cheng; Shepard, Donald B; Chong, Rebecca A; López Arriaza, José; Hall, Kathryn; Castoe, Todd A; Feschotte, Cédric; Pollock, David D; Mueller, Rachel Lockridge
Among vertebrates, most of the largest genomes are found within the salamanders, a clade of amphibians that includes 613 species. Salamander genome sizes range from ~14 to ~120 Gb. Because genome size is correlated with nucleus and cell sizes, as well as other traits, morphological evolution in salamanders has been profoundly affected by genomic gigantism. However, the molecular mechanisms driving genomic expansion in this clade remain largely unknown. Here, we present the first comparative analysis of transposable element (TE) content in salamanders. Using high-throughput sequencing, we generated genomic shotgun data for six species from the Plethodontidae, the largest family of salamanders. We then developed a pipeline to mine TE sequences from shotgun data in taxa with limited genomic resources, such as salamanders. Our summaries of overall TE abundance and diversity for each species demonstrate that TEs make up a substantial portion of salamander genomes, and that all of the major known types of TEs are represented in salamanders. The most abundant TE superfamilies found in the genomes of our six focal species are similar, despite substantial variation in genome size. However, our results demonstrate a major difference between salamanders and other vertebrates: salamander genomes contain much larger amounts of long terminal repeat (LTR) retrotransposons, primarily Ty3/gypsy elements. Thus, the extreme increase in genome size that occurred in salamanders was likely accompanied by a shift in TE landscape. These results suggest that increased proliferation of LTR retrotransposons was a major molecular mechanism contributing to genomic expansion in salamanders.
Grant, E.H.C.; Green, L.E.; Lowe, W.H.
1. Stream ecosystems exhibit a highly consistent dendritic geometry in which linear habitat units intersect to create a hierarchical network of connected branches. 2. Ecological and life history traits of species living in streams, such as the potential for overland movement, may interact with this architecture to shape patterns of occupancy and response to disturbance. Specifically, large-scale habitat alteration that fragments stream networks and reduces connectivity may reduce the probability a stream is occupied by sensitive species, such as stream salamanders. 3. We collected habitat occupancy data on four species of stream salamanders in first-order (i.e. headwater) streams in undeveloped and urbanised regions of the eastern U.S.A. We then used an information-theoretic approach to test alternative models of salamander occupancy based on a priori predictions of the effects of network configuration, region and salamander life history. 4. Across all four species, we found that streams connected to other first-order streams had higher occupancy than those flowing directly into larger streams and rivers. For three of the four species, occupancy was lower in the urbanised region than in the undeveloped region. 5. These results demonstrate that the spatial configuration of stream networks within protected areas affects the occurrences of stream salamander species. We strongly encourage preservation of network connections between first-order streams in conservation planning and management decisions that may affect stream species.
Struck Torsten H
Full Text Available Abstract Background The retention of ancestral juvenile characters by adult stages of descendants is called paedomorphosis. However, this process can mislead phylogenetic analyses based on morphological data, even in combination with molecular data, because the assessment if a character is primary absent or secondary lost is difficult. Thus, the detection of incongruence between morphological and molecular data is necessary to investigate the reliability of simultaneous analyses. Different methods have been proposed to detect data congruence or incongruence. Five of them (PABA, PBS, NDI, LILD, DRI are used herein to assess incongruence between morphological and molecular data in a case study addressing salamander phylogeny, which comprises several supposedly paedomorphic taxa. Therefore, previously published data sets were compiled herein. Furthermore, two strategies ameliorating effects of paedomorphosis on phylogenetic studies were tested herein using a statistical rigor. Additionally, efficiency of the different methods to assess incongruence was analyzed using this empirical data set. Finally, a test statistic is presented for all these methods except DRI. Results The addition of morphological data to molecular data results in both different positions of three of the four paedomorphic taxa and strong incongruence, but treating the morphological data using different strategies ameliorating the negative impact of paedomorphosis revokes these changes and minimizes the conflict. Of these strategies the strategy to just exclude paedomorphic character traits seem to be most beneficial. Of the three molecular partitions analyzed herein the RAG1 partition seems to be the most suitable to resolve deep salamander phylogeny. The rRNA and mtDNA partition are either too conserved or too variable, respectively. Of the different methods to detect incongruence, the NDI and PABA approaches are more conservative in the indication of incongruence than LILD and
Full Text Available Antimicrobial peptides have been widely identified from amphibian skins except salamanders. A novel antimicrobial peptide (CFBD was isolated and characterized from skin secretions of the salamander, Cynops fudingensis. The cDNA encoding CFBD precursor was cloned from the skin cDNA library of C. fudingensis. The precursor was composed of three domains: signal peptide of 17 residues, mature peptide of 41 residues and intervening propeptide of 3 residues. There are six cysteines in the sequence of mature CFBD peptide, which possibly form three disulfide-bridges. CFBD showed antimicrobial activities against Staphylococcus aureus, Bacillus subtilis, Candida albicans and Escherichia coli. This peptide could be classified into family of β-defensin based on its sequence similarity with β-defensins from other vertebrates. Evolution analysis indicated that CFBD was close to fish β-defensin. As far as we know, CFBD is the first β-defensin antimicrobial peptide from salamanders.
Bonett, Ronald M; Chippindale, Paul T; Moler, Paul E; Van Devender, R Wayne; Wake, David B
The Amphiumidae contains three species of elongate, permanently aquatic salamanders with four diminutive limbs that append one, two, or three toes. Two of the species, Amphiuma means and A. tridactylum, are among the largest salamanders in the world, reaching lengths of more than one meter, whereas the third species (A. pholeter), extinct amphiumids, and closely related salamander families are relatively small. Amphiuma means and A. tridactylum are widespread species and live in a wide range of lowland aquatic habitats on the Coastal Plain of the southeastern United States, whereas A. pholeter is restricted to very specialized organic muck habitats and is syntopic with A. means. Here we present analyses of sequences of mitochondrial and nuclear loci from across the distribution of the three taxa to assess lineage diversity, relationships, and relative timing of divergence in amphiumid salamanders. In addition we analyze the evolution of gigantism in the clade. Our analyses indicate three lineages that have diverged since the late Miocene, that correspond to the three currently recognized species, but the two gigantic species are not each other's closest relatives. Given that the most closely related salamander families and fossil amphiumids from the Upper Cretaceous and Paleocene are relatively small, our results suggest at least two extreme changes in body size within the Amphuimidae. Gigantic body size either evolved once as the ancestral condition of modern amphiumas, with a subsequent strong size reduction in A. pholeter, or gigantism independently evolved twice in the modern species, A. means and A. tridactylum. These patterns are concordant with differences in habitat breadth and range size among lineages, and have implications for reproductive isolation and diversification of amphiumid salamanders.
Ronald M Bonett
Full Text Available The Amphiumidae contains three species of elongate, permanently aquatic salamanders with four diminutive limbs that append one, two, or three toes. Two of the species, Amphiuma means and A. tridactylum, are among the largest salamanders in the world, reaching lengths of more than one meter, whereas the third species (A. pholeter, extinct amphiumids, and closely related salamander families are relatively small. Amphiuma means and A. tridactylum are widespread species and live in a wide range of lowland aquatic habitats on the Coastal Plain of the southeastern United States, whereas A. pholeter is restricted to very specialized organic muck habitats and is syntopic with A. means. Here we present analyses of sequences of mitochondrial and nuclear loci from across the distribution of the three taxa to assess lineage diversity, relationships, and relative timing of divergence in amphiumid salamanders. In addition we analyze the evolution of gigantism in the clade. Our analyses indicate three lineages that have diverged since the late Miocene, that correspond to the three currently recognized species, but the two gigantic species are not each other's closest relatives. Given that the most closely related salamander families and fossil amphiumids from the Upper Cretaceous and Paleocene are relatively small, our results suggest at least two extreme changes in body size within the Amphuimidae. Gigantic body size either evolved once as the ancestral condition of modern amphiumas, with a subsequent strong size reduction in A. pholeter, or gigantism independently evolved twice in the modern species, A. means and A. tridactylum. These patterns are concordant with differences in habitat breadth and range size among lineages, and have implications for reproductive isolation and diversification of amphiumid salamanders.
Burke, E.M.; Pough, F.H.
We measured temperature resistance and lactic acid production of Notophthalmus viridescens and Plethodon cinerus acclimated to 10, 20 or 30/sup 0/C. Fatigue is responsible for the Loss of Righting Response (LRR) observed when Plethodon are heated and contributes to the LRR of Notophthalmus efts. LRR is purely a manifestation of incapacitation by high temperature in Notophthalmus adults. The Critical Thermal Maximum (CTM) is solely a product of temperature incapacitation in all three forms. The standard test for temperature resistance should be expanded to include tests of fatigue.
Madison, D. M.
A detailed description is given of experiments made to determine the senses and chemical cues used by salamanders for homing orientation. Sensory impairment and cue manipulative techniques were used in the investigation. All experiments were carried out at night. Results show that sense impaired animals did not home as readily as those who were blind but retained their sensory mechanism. This fact suggests that the olfactory mechanism is necessary for homing in the salamander. It was determined that after the impaired salamander regenerated its sensory mechanism it too returned home. It was concluded that homing ability in salamanders is direction independent, distant dependent, and vision independent.
Ryan, Maureen E; Johnson, Jarrett R; Fitzpatrick, Benjamin M; Lowenstine, Linda J; Picco, Angela M; Shaffer, H Bradley
Biological invasions and habitat alteration are often detrimental to native species, but their interactions are difficult to predict. Interbreeding between native and introduced species generates novel genotypes and phenotypes, and human land use alters habitat structure and chemistry. Both invasions and habitat alteration create new biological challenges and opportunities. In the intensively farmed Salinas Valley, California (U.S.A.), threatened California tiger salamanders (Ambystoma californiense) have been replaced by hybrids between California tiger salamander and introduced barred tiger salamanders (Ambystoma tigrinum mavortium). We conducted an enclosure experiment to examine the effects habitat modification and relative frequency of hybrid and native California tiger salamanders have on recruitment of salamanders and their prey, Pacific chorus frogs (Pseudacris regilla). We tested whether recruitment differed among genetic classes of tiger salamanders (hybrid or native) and pond hydroperiod (seasonal or perennial). Roughly 6 weeks into the experiment, 70% (of 378 total) of salamander larvae died in 4 out of 6 ponds. Native salamanders survived (n = 12) in these ponds only if they had metamorphosed prior to the die-offs. During die-offs, all larvae of native salamanders died, whereas 56% of hybrid larvae died. We necropsied native and hybrid salamanders, tested water quality, and queried the California Department of Pesticide Regulation database to investigate possible causes of the die-offs. Salamander die-offs, changes in the abundance of other community members (invertebrates, algae, and cyanobacteria), shifts in salamander sex ratio, and patterns of pesticide application in adjacent fields suggest that pesticide use may have contributed to die-offs. That all survivors were hybrids suggests that environmental stress may promote rapid displacement of native genotypes.
Glorioso, Brad M.; Waddle, J. Hardin; Hefner, Jeromi M.
Reproductive aspects of Ambystoma salamanders were investigated at sites in Louisiana (2010–12) and Mississippi (2013). Three species occurred at the Louisiana site, Spotted Salamander (A. maculatum), Marbled Salamander (A. opacum), and Mole Salamander (A. talpoideum), whereas only Spotted Salamanders were studied at the Mississippi site. A total of 162 and 71 egg masses of Spotted Salamanders were examined at the Louisiana and Mississippi sites, respectively. Significantly more Spotted Salamander eggs per egg mass were observed at the Mississippi site (x̄ = 78.2) than the Louisiana site (x̄ = 53.8; P Salamanders at the Mississippi site (82.9 mm) was significantly larger than the Louisiana site (76.1 mm; P Salamander egg masses were not found at the Mississippi site, but accounted for 11% of examined egg masses at the Louisiana site. The mean number of eggs per egg mass at the Louisiana site did not differ between opaque (47.3) and clear (54.6) egg masses (P = 0.21). A total of 47 egg masses of the Mole Salamander were examined, with a mean number of 6.7 embryos per mass. Twenty-three individual nests of the Marbled Salamander were found either under or in decaying logs in the dry pond basins. There was no difference between the mean numbers of eggs per mass of attended nests (93.0) versus those that were discovered unattended (86.6; P = 0.67). Females tended to place their nests at intermediate heights within the pond basin.
Wang, Heng; Simon, András
Salamanders can regenerate entire limbs throughout their life. A critical step during limb regeneration is formation of a blastema, which gives rise to the new extremity. Salamander limb regeneration has historically been tightly linked to the term dedifferentiation, however, with refined research tools it is important to revisit the definition of dedifferentiation in the context. To what extent do differentiated cells revert their differentiated phenotypes? To what extent do progeny from differentiated cells cross lineage boundaries during regeneration? How do cell cycle plasticity and lineage plasticity relate to each other? What is the relationship between dedifferentiation of specialized cells and activation of tissue resident stem cells in terms of their contribution to the new limb? Here we highlight these problems through the case of skeletal muscle.
Madison, Dale M; Sullivan, Aaron M; Maerz, John C; McDarby, James H; Rohr, Jason R
Prey species show diverse antipredator responses to chemical cues signaling predation threat. Among terrestrial vertebrates, the red-backed salamander, Plethodon cinereus, is an important species in the study of these chemical defenses. During the day and early evening, this species avoids rinses from garter snakes, Thamnophis sirtalis, independent of snake diet, but late at night. avoids only those rinses from garter snakes that have recently eaten P. cinereus. We tested whether the selective, late-night response requires the ingestion or injury of salamanders. In three experiments, we tested P. cinereus for their responses to separate or combined rinses from salamanders (undisturbed, distressed, and injured P. cinereus) and snakes (unfed, earthworm fed, and salamander-fed T. sirtalis). When paired against a water control, only rinses from salamander-fed snakes were avoided. When salamander treatments (undisturbed or distressed) were combined with the snake treatments (unfed or earthworm-fed) and tested against a water control, the combinations elicited avoidance. When selected treatments were paired against the standard rinse from salamander-fed snakes, only the combined rinses from salamanders and snakes nullified the avoidance response to the standard rinse. These data reveal a prey defense mechanism involving chemical elements fromboth the predatorand prey that does not require injury or ingestion of the prey in the formation of the cue.
Beck, Scott; Moorman, Christopher; DePerno, Christopher S.; Simons, Theodore R.
Herpetofauna have declined globally, and monitoring is a useful approach to document local and long-term changes. However, monitoring efforts often fail to account for detectability or follow standardized protocols. We performed a case study at Hemlock Bluffs Nature Preserve in Cary, NC to model occupancy of focal species and demonstrate a replicable long-term protocol useful to parks and nature preserves. From March 2010 to 2011, we documented occupancy of Ambystoma opacum(Marbled Salamander), Plethodon cinereus (Red-backed Salamander), Carphophis amoenus (Eastern Worm Snake), and Diadophis punctatus (Ringneck Snake) at coverboard sites and estimated breeding female Ambystoma maculatum (Spotted Salamander) abundance via dependent double-observer egg-mass counts in ephemeral pools. Temperature influenced detection of both Marbled and Red-backed Salamanders. Based on egg-mass data, we estimated Spotted Salamander abundance to be between 21 and 44 breeding females. We detected 43 of 53 previously documented herpetofauna species. Our approach demonstrates a monitoring protocol that accounts for factors that influence species detection and is replicable by parks or nature preserves with limited resources.
Full Text Available Bromeliad phytotelmata are frequently used by several Neotropical amphibian taxa, possibly due to their high humidity, microclimatic stability, and role as a refuge from predators. Indeed, the ability of phytotelmata to buffer against adverse environmental conditions may be instrumental in allowing some amphibian species to survive during periods of environmental change or to colonize sub-optimal habitats. Association between bromeliad traits and salamanders has not been studied at a fine scale, despite the intimate association of many salamander species with bromeliads. Here, we identify microhabitat characteristics of epiphytic bromeliads used by two species of the Bolitoglossa morio group (B. morio and B. pacaya in forest disturbed by volcanic activity in Guatemala. Specifically, we measured multiple variables for bromeliads (height and position in tree, phytotelma water temperature and pH, canopy cover, phytotelma size, leaf size, and tree diameter at breast height, as well as salamander size. We employed a DNA barcoding approach to identify salamanders. We found that B. morio and B. pacaya occurred in microsympatry in bromeliads and that phytotelmata size and temperature of bromeliad microhabitat were the most important factors associated with the presence of salamanders. Moreover, phytotelmata with higher pH contained larger salamanders, suggesting that larger salamanders or aggregated individuals might modify pH. These results show that bromeliad selection is nonrandom with respect to microhabitat characteristics, and provide insight into the relationship between salamanders and this unique arboreal environment.
Ruano-Fajardo, Gustavo; Rovito, Sean M.; Ladle, Richard J.
Bromeliad phytotelmata are frequently used by several Neotropical amphibian taxa, possibly due to their high humidity, microclimatic stability, and role as a refuge from predators. Indeed, the ability of phytotelmata to buffer against adverse environmental conditions may be instrumental in allowing some amphibian species to survive during periods of environmental change or to colonize sub-optimal habitats. Association between bromeliad traits and salamanders has not been studied at a fine scale, despite the intimate association of many salamander species with bromeliads. Here, we identify microhabitat characteristics of epiphytic bromeliads used by two species of the Bolitoglossa morio group (B. morio and B. pacaya) in forest disturbed by volcanic activity in Guatemala. Specifically, we measured multiple variables for bromeliads (height and position in tree, phytotelma water temperature and pH, canopy cover, phytotelma size, leaf size, and tree diameter at breast height), as well as salamander size. We employed a DNA barcoding approach to identify salamanders. We found that B. morio and B. pacaya occurred in microsympatry in bromeliads and that phytotelmata size and temperature of bromeliad microhabitat were the most important factors associated with the presence of salamanders. Moreover, phytotelmata with higher pH contained larger salamanders, suggesting that larger salamanders or aggregated individuals might modify pH. These results show that bromeliad selection is nonrandom with respect to microhabitat characteristics, and provide insight into the relationship between salamanders and this unique arboreal environment. PMID:24892414
Hoffman, R.; Pearl, C.A.; Larson, G.L.; Samora, B.
Oviposition timing, behaviors, and microhabitats of ambystomatid salamanders vary considerably (Egan and Paton 2004; Figiel and Semlitsch 1995; Howard and Wallace 1985; Mac-Cracken 2007). Regardless of species, however, females typically oviposit using sites conducive to embryo development and survival. For example, the results of an experiment by Figiel and Semlitsch (1995) on Ambystoma opacum (Marbled Salamander) oviposition indicated that females actively selected sites that were under grass clumps in wet versus dry treatments, and surmised that environmental conditions such as humidity, moisture, and temperature contributed to their results. Other factors associated with ambystomatid oviposition and embryo survival include water temperature (Anderson 1972; Brown 1976), dissolved oxygen concentration (Petranka et al. 1982; Sacerdote and King 2009), oviposition depth (Dougherty et al. 2005; Egan and Paton 2004), and oviposition attachment structures such as woody vegetation (McCracken 2007; Nussbaum et al. 1983). Resetarits (1996), in creating a model of oviposition site selection for anuran amphibians, hypothesized that oviparous organisms were also capable of modifying oviposition behavior and site selection to accommodate varying habitat conditions and to minimize potential negative effects of environmental stressors. Kats and Sih (1992), investigating the oviposition of Ambystoma barbouri (Streamside Salamander) in pools of a Kentucky stream, found that females preferred pools without predatory Lepomis cyanellus (Green Sunfish), and that the number of egg masses present in a pool historically containing fish increased significantly the year after fish had been extirpated from the pool. Palen et al. (2005) determined that Ambystoma gracile (Northwestern Salamander) and Ambystoma macrodactylum (Longtoed Salamander) eggs were deposited either at increased depth or in full shaded habitats, respectively, as water transperancy to UV-B radiation increased.
Grant, Evan H. Campbell; Wofford, John E.B.; Smith, D.R.; Dennis, J.; Hawkins-Hoffman, C.; Schaberl, J.; Foley, M.; Bogle, M.
Here we report on a structured decision making (SDM) process to identify management strategies to ensure persistence of the federally endangered Shenandoah salamander (Plethodon shenandoah), given that it may be at increased extinction risk under projected climate change. The focus of this report is the second of two SDM workshops; in the first workshop, participants developed a prototype of the decision, including problem frame, management objectives and a suite of potential management strategies, predictive models to inform the decision and link alternatives with the objectives to identify potential solutions, and identified data needs to reduce key uncertainties in the decision. Participants in this second workshop included experts in National Park Service policy at multiple administrative levels, who refined objectives, further evaluated the initial management alternatives, and discussed policy constraints on implementing active management for the species and its high-elevation habitat. The conclusion of the second workshop was similar to that of the first: the current state of information and objectives suggest that there is some value in considering active management to reduce the long-term extinction risk for the species, though there are institutional conservative policies to implementing active management at range-wide scales. The workshop participants also emphasized a conservative NPS management philosophy, including caution in implementing management actions that may ultimately harm the system, a stated assumption that ecosystem changes were “natural” unless demonstrated otherwise (therefore not warranting active management to mitigate), and a need to demonstrate that extinction risk is tied to anthropogenic influence prior to taking active management to mitigate specific anthropogenic influences. Even within a protected area having minimal human disturbance, intertwined environmental variables and interspecific relationships that drive population
Nemecek, Julie; Nag, Nabanita; Carlson, Christina M.; Schneider, Jay R.; Heisey, Dennis M.; Johnson, Christopher J.; Asher, David M.; Gregori, Luisa
Rapid antemortem tests to detect individuals with transmissible spongiform encephalopathies (TSE) would contribute to public health. We investigated a technique known as protein misfolding cyclic amplification (PMCA) to amplify abnormal prion protein (PrPTSE) from highly diluted variant Creutzfeldt-Jakob disease (vCJD)-infected human and macaque brain homogenates, seeking to improve the rapid detection of PrPTSE in tissues and blood. Macaque vCJD PrPTSE did not amplify using normal macaque brain homogenate as substrate (intraspecies PMCA). Next, we tested interspecies PMCA with normal brain homogenate of the southern red-backed vole (RBV), a close relative of the bank vole, seeded with macaque vCJD PrPTSE. The RBV has a natural polymorphism at residue 170 of the PrP-encoding gene (N/N, S/S, and S/N). We investigated the effect of this polymorphism on amplification of human and macaque vCJD PrPTSE. Meadow vole brain (170N/N PrP genotype) was also included in the panel of substrates tested. Both humans and macaques have the same 170S/S PrP genotype. Macaque PrPTSE was best amplified with RBV 170S/S brain, although 170N/N and 170S/N were also competent substrates, while meadow vole brain was a poor substrate. In contrast, human PrPTSE demonstrated a striking narrow selectivity for PMCA substrate and was successfully amplified only with RBV 170S/S brain. These observations suggest that macaque PrPTSE was more permissive than human PrPTSE in selecting the competent RBV substrate. RBV 170S/S brain was used to assess the sensitivity of PMCA with PrPTSE from brains of humans and macaques with vCJD. PrPTSE signals were reproducibly detected by Western blot in dilutions through 10-12 of vCJD-infected 10% brain homogenates. This is the first report showing PrPTSE from vCJD-infected human and macaque brains efficiently amplified with RBV brain as the substrate. Based on our estimates, PMCA showed a sensitivity that might be sufficient to detect PrPTSE in v
Full Text Available Rapid antemortem tests to detect individuals with transmissible spongiform encephalopathies (TSE would contribute to public health. We investigated a technique known as protein misfolding cyclic amplification (PMCA to amplify abnormal prion protein (PrP(TSE from highly diluted variant Creutzfeldt-Jakob disease (vCJD-infected human and macaque brain homogenates, seeking to improve the rapid detection of PrP(TSE in tissues and blood. Macaque vCJD PrP(TSE did not amplify using normal macaque brain homogenate as substrate (intraspecies PMCA. Next, we tested interspecies PMCA with normal brain homogenate of the southern red-backed vole (RBV, a close relative of the bank vole, seeded with macaque vCJD PrP(TSE. The RBV has a natural polymorphism at residue 170 of the PrP-encoding gene (N/N, S/S, and S/N. We investigated the effect of this polymorphism on amplification of human and macaque vCJD PrP(TSE. Meadow vole brain (170N/N PrP genotype was also included in the panel of substrates tested. Both humans and macaques have the same 170S/S PrP genotype. Macaque PrP(TSE was best amplified with RBV 170S/S brain, although 170N/N and 170S/N were also competent substrates, while meadow vole brain was a poor substrate. In contrast, human PrP(TSE demonstrated a striking narrow selectivity for PMCA substrate and was successfully amplified only with RBV 170S/S brain. These observations suggest that macaque PrP(TSE was more permissive than human PrP(TSE in selecting the competent RBV substrate. RBV 170S/S brain was used to assess the sensitivity of PMCA with PrP(TSE from brains of humans and macaques with vCJD. PrP(TSE signals were reproducibly detected by Western blot in dilutions through 10⁻¹² of vCJD-infected 10% brain homogenates. This is the first report showing PrP(TSE from vCJD-infected human and macaque brains efficiently amplified with RBV brain as the substrate. Based on our estimates, PMCA showed a sensitivity that might be sufficient to detect Pr
Price, Steven J; Cecala, Kristen K; Browne, Robert A; Dorcas, Michael E
Urban development is the most common form of land conversion in the United States. Using a before-after control-impact study design, we investigated the effects of urbanization on larval and adult stages of southern two-lined salamanders (Eurycea cirrigera) and northern dusky salamanders (Desmognathus fuscus). Over 5 years, we estimated changes in occupancy and probabilities of colonization and survival in 13 stream catchments after urbanization and in 17 catchments that were not urbanized. We also examined effects of proportion of urbanized area in a catchment and distance of the salamander population to the nearest stream on probabilities of colonization and survival. Before urbanization, adult and larval stages of the two salamander species occupied nearly all surveyed streams, with occupancy estimates ranging from 1.0 to 0.78. Four years after urbanization mean occupancy of larval and adult two-lined salamanders had decreased from 0.87 and 0.78 to 0.57 and 0.39, respectively. Estimates of mean occupancy of larval northern dusky salamanders decreased from 1.0 to 0.57 in urban streams 4 years after urbanization; however, adult northern dusky salamander occupancy remained close to 1.0 in urban streams over 5 years. Occupancy estimates in control streams were similar for each species and stage over 5 years. Urbanization was associated with decreases in survival probabilities of adult and larval two-lined salamanders and decreases in colonization probabilities of larval dusky salamanders. Nevertheless, proportion of impervious surface and distance to nearest stream had little effect on probabilities of survival and colonization. Our results imply that in the evaluation of the effects of urbanization on species, such as amphibians, with complex life cycles, consideration of the effects of urbanization on both adult and larval stages is required.
Ijspeert, Auke J.; Arbib, Michael A.
This article investigates the neural mechanisms underlying locomotion and visually-guided behavior in a lower vertebrate: the salamander. We develop connectionist models of the salamander's locomotor circuitry and visual system, and analyze their functioning by embedding them into a biomechanical simulation of the salamander's body. This work is therefore an experiment in computational neuroethology which aims at investigating how behavior results from the coupling of a central nervous system (CNS) and a body, and from the interactions of the CNS-body pair with the environment. We believe that understanding these mechanisms is not only relevant for neurobiology but also for potential applications in robotics.
Christensen, Christian Bech; Lauridsen, Henrik; Christensen-Dalsgaard, Jakob
hearing in early tetrapods. Here, we combine imaging techniques with neurophysiological measurements to resolve how the change from aquatic larvae to terrestrial adult affects the ear morphology and sensory capabilities of salamanders. We show that air-induced pressure detection enhances underwater...... hearing sensitivity of salamanders at frequencies above 120 Hz, and that both terrestrial adults and fully aquatic juvenile salamanders can detect airborne sound. Collectively, these findings suggest that early atympanic tetrapods may have been pre-equipped to aerial hearing and are able to hear airborne...
Kumar, Anoop; Delgado, Jean Paul
Limb regeneration of salamanders is nerve dependent, and the removal of the nerves in early stages of limb regeneration severely curtails the proliferation of the blastemal cells and growth of the regenerate. The removal of the neural tube from a developing salamander embryo results in an aneurogenic larva and the aneurogenic limb (ANL) develops independently without innervation. Paradoxically, the limb in an ANL is capable of regeneration in a nerve-independent manner. Here, we describe a detailed method for the generation of ANL in the spotted salamander, Ambystoma maculatum, for regeneration studies.
Wynn, Addison H
A contact zone between two electrophoretically differentiated forms of Plethodon cinereus extends across the Del-Mar-Va Peninsula. Based on the presumed nature of regional climates during the late Pleistocene, it is suggested that the climate was inhospitable for P. cinereus on the Del-Mar-Va Peninsula at the Wisconsin maximum. It is postulated that, after the Laurentide ice sheet receded about 14,000 years BP, the peninsula was invaded by populations of P. cinereus from the north and south. When the two forms of P. cinereus met, a secondary zone of contact was produced. This contact zone is now apparent in an area in which six electrophoretic loci have concordant clinal changes in allele frequency. The lack of evidence for heterozygote deficiencies, the broad width of the contact zone, and the overall symmetry of introgression suggest that fusion of the populations is occurring freely. Two esterase loci are in linkage disequilibrium in four of eight populations to the north of the contact zone. As all populations of P. cinereus studied to date have disequilibrium coefficients of similar sign, selection is likely important in the maintenance of this linkage disequilibrium. Within the contact zone, the incidence of the linkage disequilibrium increases, but exchange of genes between the two introgressing populations may account for the higher incidence of linkage disequilibrium. © 1986 The Society for the Study of Evolution.
Dinsmore, C E
As an epimorphic system, the urodele tail has much in common with the urodele limb, relative to tissue components. In an effort to elucidate potentially similar functions of the skin during regeneration, two procedures which have been shown to be highly disruptive of limb morphogenesis were performed on the tail. In the first, cuffs of tail skin were rotated 180 degrees about the long axis of tails in Plethodon cinereus. Subsequent amputation through the rotated cuffs produced regenerates at a normal rate which were also normal in both internal architecture and skin gland distribution. In a second series, cuffs of tail skin were rotated 90 degrees about the dorso-ventral axis of the tail, such that the dorsal red stripe encircled the tail. Upon amputation through the middle of the cuff, the stump skin presented an exclusively dorsal surface which, nevertheless, had no effect upon the morphogenetic success of the regenerate. These data therefore indicate that a tissue, such as skin, may be inconsistent in its morphogenetic influence among epimorphic fields within the same organism. It is not clear whether these results reflect differences in field behavior or in absolute tissue potential determined by the field of origin. A tissue hierarchy of morphogenetic influence within and among epimorphic fields is suggested as a preliminary framework within which to coordinate such information.
Reinhard, Sandy; Renner, Sandra; Kupfer, Alexander
We analysed sexual size dimorphism (SSD) for two Mediterranean species of the "true" salamander clade possessing distinct life histories (Salamandra algira and Mertensiella caucasica) and equilibrated the morphometric approach to individual age by using skeletochronology. For species that have a short breeding season and live at high altitudes, such as Mediterranean amphibians, the fecundity advantage hypothesis predicts female-biased SSD to maximise reproductive success. Our results showed no SSD in either species; however, morphometric data indicated a male-biased dimorphism in limb (arm and leg) dimensions in both species when compared to body size. Limb dimorphisms are likely related to the particular mating system, which involves an amplexus during spermatophore transfer. Arm length appeared sexually dimorphic during ontogeny both in viviparous S. algira and oviparous M. caucasica. A review on SSD indicated monomorphy of body size as a common lineage-specific pattern among the "true" salamander clade, but also the common presence of other traits such as sexually dimorphic limb proportions.
US Fish and Wildlife Service, Department of the Interior — The objectives of this project are to (1) conduct transect and quadrat sampling for streamside salamanders, (2) determine detection rates and population estimates...
US Fish and Wildlife Service, Department of the Interior — The objectives of this project are to (1) conduct transect and quadrat sampling for streamside salamanders, (2) determine detection rates and population estimates...
US Fish and Wildlife Service, Department of the Interior — The objectives of this project are to (1) conduct transect and quadrat sampling for streamside salamanders, (2) determine detection rates and population estimates...
US Fish and Wildlife Service, Department of the Interior — These data identify, in general, the areas of final critical habitat for the endangered Ambystoma bishopi (reticulated flatwoods salamander).
US Fish and Wildlife Service, Department of the Interior — These data identify, in general, the areas of final critical habitat for the endangered Ambystoma bishopi (reticulated flatwoods salamander).
US Fish and Wildlife Service, Department of the Interior — The primary goal for this project is to establish baseline information on populations of the Cheat Mountain salamander on the refuge. In the future, an additional...
US Fish and Wildlife Service, Department of the Interior — To provide the user with a general idea of areas where final critical habitat for San Marcos salamander (Eurycea nana) occur based on the description provided in the...
Ramotnik, Cindy A.; Scott, N.J.
We measured habitat components for two state-listed endangered salamanders in New Mexico in 1986 and 1987. Both species are restricted to mesic environments within high-elevation, mixed coniferous forests. Steep slope and high elevation were the most useful variables for predicting the occurrence of Jemez Mountains salamanders and Sacramento Mountain salamanders, respectively. Although the discriminant models show some predictive value in detecting salamanders based on habitat variables, we believe that the best survey technique is ground-truth surveys in wet weather. A better fit of the discriminant models might be obtained by including variables not measured e.g., fire and logging history, and soil characteristics. We offer interim management guidelines as a result of our analysis.
Sutherland, Chris; Munoz, David; Miller, David A.W.; Grant, Evan
Spatial capture–recapture (SCR) is a relatively recent development in ecological statistics that provides a spatial context for estimating abundance and space use patterns, and improves inference about absolute population density. SCR has been applied to individual encounter data collected noninvasively using methods such as camera traps, hair snares, and scat surveys. Despite the widespread use of capture-based surveys to monitor amphibians and reptiles, there are few applications of SCR in the herpetological literature. We demonstrate the utility of the application of SCR for studies of reptiles and amphibians by analyzing capture–recapture data from Red-Backed Salamanders, Plethodon cinereus, collected using artificial cover boards. Using SCR to analyze spatial encounter histories of marked individuals, we found evidence that density differed little among four sites within the same forest (on average, 1.59 salamanders/m2) and that salamander detection probability peaked in early October (Julian day 278) reflecting expected surface activity patterns of the species. The spatial scale of detectability, a measure of space use, indicates that the home range size for this population of Red-Backed Salamanders in autumn was 16.89 m2. Surveying reptiles and amphibians using artificial cover boards regularly generates spatial encounter history data of known individuals, which can readily be analyzed using SCR methods, providing estimates of absolute density and inference about the spatial scale of habitat use.
Arenas, Claudia Marcela; Gómez-Molina, Andrea; Delgado, Jean Paul
Limb regeneration studies have been extensively carried out in species of Ambystomatidae and Salamandridae families. So far limited research has been conducted in species belonging to the Plethodontidae family, where some of the species differs from other salamander families due to their direct development, thus absence of a larval life. Here, we describe a protocol to maintain the plethodontid salamanders of genus Bolitoglossa species under laboratory conditions to perform regeneration studies.
Wood, Petra Bohall; Williams, Jennifer M.
Mountaintop removal mining, a large-scale disturbance affecting vegetation, soil structure, and topography, converts landscapes from mature forests to extensive grassland and shrubland habitats. We sampled salamanders using drift-fence arrays and coverboard transects on and near mountaintop removal mines in southern West Virginia, USA, during 2000–2002. We compared terrestrial salamander relative abundance and species richness of un-mined, intact forest with habitats on reclaimed mountaintop removal mines (reclaimed grassland, reclaimed shrubland, and fragmented forest). Salamanders within forests increased in relative abundance with increasing distance from reclaimed mine edge. Reclaimed grassland and shrubland habitats had lower relative abundance and species richness than forests. Characteristics of reclaimed habitats that likely contributed to lower salamander abundance included poor soils (dry, compacted, little organic matter, high rock content), reduced vertical structure of vegetation and little tree cover, and low litter and woody debris cover. Past research has shown that salamander populations reduced by clearcutting may rebound in 15–24 years. Time since disturbance was 7–28 years in reclaimed habitats on our study areas and salamander populations had not reached levels found in adjacent mature forests.
Sun, Cheng; Mueller, Rachel Lockridge
Among animals, genome sizes range from 20 Mb to 130 Gb, with 380-fold variation across vertebrates. Most of the largest vertebrate genomes are found in salamanders, an amphibian clade of 660 species. Thus, salamanders are an important system for studying causes and consequences of genomic gigantism. Previously, we showed that plethodontid salamander genomes accumulate higher levels of long terminal repeat (LTR) retrotransposons than do other vertebrates, although the evolutionary origins of such sequences remained unexplored. We also showed that some salamanders in the family Plethodontidae have relatively slow rates of DNA loss through small insertions and deletions. Here, we present new data from Cryptobranchus alleganiensis, the hellbender. Cryptobranchus and Plethodontidae span the basal phylogenetic split within salamanders; thus, analyses incorporating these taxa can shed light on the genome of the ancestral crown salamander lineage, which underwent expansion. We show that high levels of LTR retrotransposons likely characterize all crown salamanders, suggesting that disproportionate expansion of this transposable element (TE) class contributed to genomic expansion. Phylogenetic and age distribution analyses of salamander LTR retrotransposons indicate that salamanders' high TE levels reflect persistence and diversification of ancestral TEs rather than horizontal transfer events. Finally, we show that relatively slow DNA loss rates through small indels likely characterize all crown salamanders, suggesting that a decreased DNA loss rate contributed to genomic expansion at the clade's base. Our identification of shared genomic features across phylogenetically distant salamanders is a first step toward identifying the evolutionary processes underlying accumulation and persistence of high levels of repetitive sequence in salamander genomes.
... Salamander, Calaveras County, CA AGENCY: Fish and Wildlife Service, Interior. ACTION: Notice of availability... salamander (tiger salamander). The applicant would implement a conservation program to minimize and mitigate... comment is in reference to the Low-Effect Habitat Conservation Plan for the California Tiger...
Brockes, Jeremy P
Regeneration is studied in a few model species of salamanders, but the ten families of salamanders show considerable variation, and this has implications for our understanding of salamander biology. The most recent classification of the families identifies the cryptobranchoidea as the basal group which diverged in the early Jurassic. Variation in the sizes of genomes is particularly obvious, and reflects a major contribution from transposable elements which is already present in the basal group.Limb development has been a focus for evodevo studies, in part because of the variable property of pre-axial dominance which distinguishes salamanders from other tetrapods. This is thought to reflect the selective pressures that operate on a free-living aquatic larva, and might also be relevant for the evolution of limb regeneration. Recent fossil evidence suggests that both pre-axial dominance and limb regeneration were present 300 million years ago in larval temnospondyl amphibians that lived in mountain lakes. A satisfying account of regeneration in salamanders may need to address all these different aspects in the future.
Mu, Lixian; Tang, Jing; Liu, Han; Shen, Chuanbin; Rong, Mingqiang; Zhang, Zhiye; Lai, Ren
Although it is well known that wound healing proceeds incredibly quickly in urodele amphibians, such as newts and salamanders, little is known about skin-wound healing, and no bioactive/effector substance that contributes to wound healing has been identified from these animals. As a step toward understanding salamander wound healing and skin regeneration, a potential wound-healing-promoting peptide (tylotoin; KCVRQNNKRVCK) was identified from salamander skin of Tylototriton verrucosus. It shows comparable wound-healing-promoting ability (EC50=11.14 μg/ml) with epidermal growth factor (EGF; NSDSECPLSHDGYCLHDGVCMYIEALDKYACNCVVGYIGERCQYRDLKWWELR) in a murine model of full-thickness dermal wound. Tylotoin directly enhances the motility and proliferation of keratinocytes, vascular endothelial cells, and fibroblasts, resulting in accelerated reepithelialization and granulation tissue formation in the wound site. Tylotoin also promotes the release of transforming growth factor β1 (TGF-β1) and interleukin 6 (IL-6), which are essential in the wound healing response. Gene-encoded tylotoin secreted in salamander skin is possibly an effector molecule for skin wound healing. This study may facilitate understanding of the cellular and molecular events that underlie quick wound healing in salamanders.
The morphology and organogenesis of Chinese giant salamander, Andrias davidianus, in its different developmental periods and stages are described in detail, which provides an intact criterion for distinguishing different stages of its developmental process.Based on the external morphological and internal histological features, six periods including 20 stages of organogenesis of Chinese giant salamander are established, which are cleavage period, blastula period, gastrula period, neurula period, organogenesis stage and hatching stage. Generally, the embryonic development of Chinese giant salamander is consistent with those of Eastern newt, Cynops orientalis,and Black spots frog, R. nigromaculata. However, they have some differences in the early cleavage process and the development of digestive system. The cleavage of Chinese giant salamander, A. davidianus is not a discoidal division type, which is different from other species reported. And the first three cleavages being meridional and a retardant development of its digestive system without halter and sucker existing are the evident features of the embryonic development of Chinese giant salamander.
Burke, John N; Bergeron, Christine M; Todd, Brian D; Hopkins, William A
Mercury (Hg) causes a range of deleterious effects in wildlife, but little is known about its effects on amphibians. Our objective was to determine whether Hg affects performance and behavior in two-lined salamanders (Eurycea bislineata). We collected salamanders from Hg-contaminated and reference sites and assessed speed, responsiveness, and prey capture ability. Mercury concentrations were >17× higher in salamanders from the contaminated sites and were among the highest documented in amphibians. In the first, but not in the second, locomotion trial, we found a significant effect of Hg on speed and responsiveness. In the prey capture experiment, reference salamanders ate approximately twice as many prey items as the contaminated salamanders. Together, our results suggest that sublethal Hg concentrations may negatively affect salamanders by reducing their ability to successfully execute tasks critical to survival. Future work is warranted to determine whether Hg has other sublethal effects on salamanders and whether other amphibians are similarly affected.
Connette, Grant M; Crawford, John A; Peterman, William E
An increasing number of studies have demonstrated relationships between climate trends and body size change of organisms. In many cases, climate might be expected to influence body size by altering thermoregulation, energetics or food availability. However, observed body size change can result from a variety of ecological processes (e.g. growth, selection, population dynamics) or imperfect observation of biological systems. We used two extensive datasets to evaluate alternative mechanisms for recently reported changes in the observed body size of plethodontid salamanders. We found that mean adult body size of salamanders can be highly sensitive to survey conditions, particularly rainfall. This systematic bias in the detection of larger or smaller individuals could result in a signature of body size change in relation to reported climate trends when it is simply observation error. We also identify considerable variability in body size distributions among years and find that individual growth rates can be strongly influenced by weather. Finally, our study demonstrates that measures of mean adult body size can be highly variable among surveys and that large sample sizes may be required to make reliable inferences. Identifying the effects of climate change is a critical area of research in ecology and conservation. Researchers should be aware that observed changes in certain organisms can result from multiple ecological processes or systematic bias due to nonrandom sampling of populations.
Full Text Available Global warming is influencing the biology of the world’s biota. Temperature increases are occurring at a faster pace than that experienced by organisms in their evolutionary histories, limiting the organisms’ response to new conditions. Mechanistic models that include physiological traits can help predict species’ responses to warming. Changes in metabolism at high temperatures are often examined; yet many species are behaviorally shielded from high temperatures. Salamanders generally favor cold temperatures and are one of few groups of metazoans to be most species-rich in temperate regions. I examined variation in body temperature, behavioral activity, and temperature dependence of resting heart rate, used as a proxy for standard metabolic rate, in fire salamanders (Salamandra salamandra. Over 26 years, I found that salamanders are behaviorally active at temperatures as low as 1 °C, and aestivate at temperatures above 16 °C. Infrared thermography indicates limited thermoregulation opportunities for these nocturnal amphibians. Temperature affects resting heart rate, causing metabolic depression above 11 °C, and metabolic compensation below 8 °C: heart rate at 3 °C is 224% the expected heart rate. Thus, salamanders operating at low temperatures during periods of peak behavioral activity are able to maintain a higher metabolic rate than the rate expected in absence of compensation. This compensatory mechanism has important ecological implications, because it increases estimated seasonal heart rates. Increased heart rate, and thus metabolism, will require higher caloric intake for field-active salamanders. Thus, it is important to consider a species performance breadth over the entire temperature range, and particularly low temperatures that are ecologically relevant for cold tolerant species such as salamanders.
Bryson Eric J
Full Text Available Abstract Background GABAergic inhibition and effects of intracellular chloride ions on calcium channel activity have been proposed to regulate neurotransmission from photoreceptors. To assess the impact of these and other chloride-dependent mechanisms on release from cones, the chloride equilibrium potential (ECl was determined in red-sensitive, large single cones from the tiger salamander retinal slice. Results Whole cell recordings were done using gramicidin perforated patch techniques to maintain endogenous Cl- levels. Membrane potentials were corrected for liquid junction potentials. Cone resting potentials were found to average -46 mV. To measure ECl, we applied long depolarizing steps to activate the calcium-activated chloride current (ICl(Ca and then determined the reversal potential for the current component that was inhibited by the Cl- channel blocker, niflumic acid. With this method, ECl was found to average -46 mV. In a complementary approach, we used a Cl-sensitive dye, MEQ, to measure the Cl- flux produced by depolarization with elevated concentrations of K+. The membrane potentials produced by the various high K+ solutions were measured in separate current clamp experiments. Consistent with electrophysiological experiments, MEQ fluorescence measurements indicated that ECl was below -36 mV. Conclusions The results of this study indicate that ECl is close to the dark resting potential. This will minimize the impact of chloride-dependent presynaptic mechanisms in cone terminals involving GABAa receptors, glutamate transporters and ICl(Ca.
Cabelguen, Jean-Marie; Charrier, Vanessa; Mathou, Alexia
Most investigations on tetrapod locomotion have been concerned with limb movements. However, there is compelling evidence that the axial musculoskeletal system contributes to important functions during locomotion. Adult salamanders offer a remarkable opportunity to examine these functions because these amphibians use axial undulations to propel themselves in both aquatic and terrestrial environments. In this article, we review the currently available biological data on axial functions during various locomotor modes in salamanders. We also present data showing the modular organisation of the neural networks that generate axial synergies during locomotion. The functional implication of this modular organisation is discussed.
Deban, S.M.; O'Reilly, J.C.; Dicke, U.; Leeuwen, van J.L.
Many plethodontid salamanders project their tongues ballistically at high speed and for relatively great distances. Capturing evasive prey relies on the tongue reaching the target in minimum time, therefore it is expected that power production, or the rate of energy release, is maximized during tong
Wiens, John J; Hoverman, Jason T
The loss of digits is a widespread evolutionary trend in tetrapods which occurs in nearly every major clade. Alberch and Gale showed that the order in which digits are evolutionarily lost in salamanders versus frogs corresponds to the order in which they develop in each group, providing a classic example of developmental constraint. However, what actually drives the loss of digits in salamanders has remained unclear. Alberch and Gale suggested that loss of digits might be associated with paedomorphosis or with reduced body size. We test these hypotheses by combining morphometric and phylogenetic information for 98 species of salamanders. We find that digit loss is associated with both paedomorphosis and reduction in body size. However, these trends are surprisingly contradictory, in that paedomorphosis is significantly associated with an increase in body size in salamanders. Thus, much of the extreme digit reduction is found in the smaller species within paedomorphic clades that have, on average, unusually large body size. Our results show that the consequences of changes in body size on morphology are highly context dependent. We also show (possibly for the first time) a significant association between paedomorphosis and increased body size, rather than the expected association with reduced body size.
Chen, Y.; Znoiko, S.; Grip, W.J. de; Crouch, R.K.; Ma, J.X.
The tiger salamander lives in shallow water with bright light in the aquatic phase, and in dim tunnels or caves in the terrestrial phase. In the aquatic phase, there are five types of photoreceptors--two types of rods and three types of cones. Our previous studies showed that the green rods and
Richard C. Bruce
This study investigates interspecific variation in age at first reproduction, fecundity, and body size in multispecies assemblages of desmognathine salamanders. The hypotheses tested are that interspecific differences in body size among desmognathines stem proximately from variation in age at first reproduction and that variation in the latter trait is positively...
F. Harvey Pough; Richard E. Wilson
The two species of mole salamander that occur in the Ithaca, New York, region (Ambystoma maculatum and A. jeffersonianum) breed in temporary ponds that are formed by accumulation of melted snow and spring rains. Water in many of these pools during the breeding season is acid; pH values as low as 3.5 have been measured. In...
Sun, Cheng; López Arriaza, José R; Mueller, Rachel Lockridge
Evolutionary changes in genome size result from the combined effects of mutation, natural selection, and genetic drift. Insertion and deletion mutations (indels) directly impact genome size by adding or removing sequences. Most species lose more DNA through small indels (i.e., ~1-30 bp) than they gain, which can result in genome reduction over time. Because this rate of DNA loss varies across species, small indel dynamics have been suggested to contribute to genome size evolution. Species with extremely large genomes provide interesting test cases for exploring the link between small indels and genome size; however, most large genomes remain relatively unexplored. Here, we examine rates of DNA loss in the tetrapods with the largest genomes-the salamanders. We used low-coverage genomic shotgun sequence data from four salamander species to examine patterns of insertion, deletion, and substitution in neutrally evolving non-long terminal repeat (LTR) retrotransposon sequences. For comparison, we estimated genome-wide DNA loss rates in non-LTR retrotransposon sequences from five other vertebrate genomes: Anolis carolinensis, Danio rerio, Gallus gallus, Homo sapiens, and Xenopus tropicalis. Our results show that salamanders have significantly lower rates of DNA loss than do other vertebrates. More specifically, salamanders experience lower numbers of deletions relative to insertions, and both deletions and insertions are skewed toward smaller sizes. On the basis of these patterns, we conclude that slow DNA loss contributes to genomic gigantism in salamanders. We also identify candidate molecular mechanisms underlying these differences and suggest that natural variation in indel dynamics provides a unique opportunity to study the basis of genome stability.
Basile, Marco; Romano, Antonio; Costa, Andrea; Posillico, Mario; Scinti Roger, Daniele; Crisci, Aldo; Raimondi, Ranieri; Altea, Tiziana; Garfì, Vittorio; Santopuoli, Giovanni; Marchetti, Marco; Salvidio, Sebastiano; De Cinti, Bruno; Matteucci, Giorgio
Many small terrestrial vertebrates exhibit limited spatial movement and are considerably exposed to changes in local environmental variables. Among such vertebrates, amphibians at present experience a dramatic decline due to their limited resilience to environmental change. Since the local survival and abundance of amphibians is intrinsically related to the availability of shelters, conservation plans need to take microhabitat requirements into account. In order to gain insight into the terrestrial ecology of the spectacled salamander Salamandrina perspicillata and to identify appropriate forest management strategies, we investigated the salamander's seasonal variability in habitat use of trees as shelters in relation to tree features (size, buttresses, basal holes) and environmental variables in a beech forest in Italy. We used the occupancy approach to assess tree suitability on a non-conventional spatial scale. Our approach provides fine-grained parameters of microhabitat suitability and elucidates many aspects of the salamander's terrestrial ecology . Occupancy changed with the annual life cycle and was higher in autumn than in spring, when females were found closer to the stream in the study area. Salamanders showed a seasonal pattern regarding the trees they occupied and a clear preference for trees with a larger diameter and more burrows. With respect to forest management, we suggest maintaining a suitable number of trees with a trunk diameter exceeding 30 cm. A practice of selective logging along the banks of streams could help maintain an adequate quantity of the appropriate microhabitat. Furthermore, in areas with a presence of salamanders, a good forest management plan requires leaving an adequate buffer zone around streams, which should be wider in autumn than in spring.
Among vertebrates, urodele amphibians are the only tetrapods with the ability to regenerate complex structures such as limbs, tail, and spinal cord throughout their lives. Furthermore, the salamander regeneration process has been shown to reverse tumorigenicity. Fibroblasts are essential for salamander regeneration, but the mechanisms underlying their role in the formation of a regeneration blastema remain unclear. Here, I review the role of fibroblasts in salamander limb regeneration and how...
Jéssica Barata da Silva; Pablo Suárez; Cleusa Yoshiko Nagamachi; Timothy Frederick Carter; Julio Cesar Pieczarka
Plethodontid salamanders of genus Bolitoglossa constitute the largest and most diverse group of salamanders, including around 20% of living caudate species. Recent studies have indicated the occurrence of five recognized species in the Brazilian Amazon Rainforest. We present here the first cytogenetic data of a Brazilian salamander, which may prove to be a useful by contribution to the cytotaxonomy of the genus. Specimens were collected near the “type” locality (Utinga, Belém, PA, Brazil). Ch...
Wood, Petra Bohall; Williams, Jennifer M.
Valley fills associated with mountaintop-removal mining bury stream headwaters and affect water quality and ecological function of reaches below fills. We quantified relative abundance of streamside salamanders in southern West Virginia during 2002 in three streams below valley fills (VFS) and in three reference streams (RS). We surveyed 36 10- × 2-m stream transects, once in summer and fall, paired by order and structure. Of 2,343 salamanders captured, 66.7% were from RS. Total salamanders (adults plus larvae) were more abundant in RS than VFS for first-order and second-order reaches. Adult salamanders had greater abundance in first-order reaches of RS than VFS. Larval salamanders were more abundant in second-order reaches of RS than VFS. No stream width or mesohabitat variables differed between VFS and RS. Only two cover variables differed. Silt cover, greater in VFS than RS first-order reaches, is a likely contributor to reduced abundance of salamanders in VFS. Second-order RS had more boulder cover than second-order VFS, which may have contributed to the higher total and larval salamander abundance in RS. Water chemistry assessments of our VFS and RS reported elevated levels of metal and ion concentrations in VFS, which can depress macroinvertebrate populations and likely affect salamander abundance. Valley fills appear to have significant negative effects on stream salamander abundance due to alterations in habitat structure, water quality and chemistry, and macroinvertebrate communities in streams below fills.
Rothermel, Betsie B; Travis, Emilie R; Miller, Debra L; Hill, Robert L; McGuire, Jessica L; Yabsley, Michael J
The interactive effects of environmental stressors and emerging infectious disease pose potential threats to stream salamander communities and their headwater stream ecosystems. To begin assessing these threats, we conducted occupancy surveys and pathogen screening of stream salamanders (Family Plethodontidae) in a protected southern Appalachians watershed in Georgia and North Carolina, USA. Of the 101 salamanders screened for both chytrid fungus (Batrachochytrium dendrobatidis) and Ranavirus, only two exhibited low-level chytrid infections. Prevalence of Ranavirus was much higher (30.4% among five species of Desmognathus). Despite the ubiquity of ranaviral infections, we found high probabilities of site occupancy (≥0.60) for all stream salamander species.
... Salamander, AT&T Portable Generator Storage Facility, Yolo County, CA AGENCY: Fish and Wildlife Service... potential for ``take'' of one Federally listed animal, the California tiger salamander. The applicant would... for the California tiger salamander into a new storage facility for portable generators within...
... Tiger Salamander, Smith's Blue Butterfly, and Yadon's Piperia at Palo Corona Regional Park, Monterey... federally threatened California red-legged frog (Rana draytonii) and California tiger salamander (Ambystoma..., California tiger salamander, Smith's blue butterfly, and Yadon's piperia on the property subject to...
Blankers, Thomas; Adams, D C; Wiens, J J
A major goal of evolutionary biology is to explain morphological diversity among species. Many studies suggest that much morphological variation is explained by adaptation to different microhabitats. Here, we test whether morphology and microhabitat use are related in plethodontid salamanders, which contain the majority of salamander species, and have radiated into a striking diversity of microhabitats. We obtained microhabitat data for 189 species that also had both morphometric and phylogenetic data. We then tested for associations between morphology and microhabitat categories using phylogenetic comparative methods. Associations between morphology and ecology in plethodontids are largely confined to a single clade within one subfamily (Bolitoglossinae), whereas variation in morphology across other plethodontids is unrelated to microhabitat categories. These results demonstrate that ecological radiation and morphological evolution can be largely decoupled in a major clade. The results also offer a striking contrast to lizards, which typically show close relationships between morphology and microhabitat.
Godwin, James W.; Pinto, Alexander R.; Nadia A. Rosenthal
The failure to replace damaged body parts in adult mammals results from a muted growth response and fibrotic scarring. Although infiltrating immune cells play a major role in determining the variable outcome of mammalian wound repair, little is known about the modulation of immune cell signaling in efficiently regenerating species such as the salamander, which can regrow complete body structures as adults. Here we present a comprehensive analysis of immune signaling during limb regeneration i...
Spotted salamanders breed in temporary pools formed in early spring by melted snow and rain. Many of these pools reflect the low pH of precipitation in the northeastern United States. Egg mortality is low (<1 percent) in pools near neutrality, but high (>60 percent) in pools more acid than pH 6. Developmental anomalies and the embryonic stage at which death occurs are the same in field situations as at corresponding pH's in laboratory experiments.
McLean, Kyle I.; Stockwell, Craig A.; Mushet, David M.
Barred tiger salamanders [Ambystoma mavortium (Baird, 1850)] exhibit two trophic morphologies; a typical and a cannibalistic morph. Cannibalistic morphs, distinguished by enlarged vomerine teeth, wide heads, slender bodies, and cannibalistic tendencies, are often found where conspecifics occur at high density. During 2012 and 2013, 162 North Dakota wetlands and lakes were sampled for salamanders. Fifty-one contained A. mavortium populations; four of these contained cannibalistic morph individuals. Two populations with cannibalistic morphs occurred at sites with high abundances of conspecifics. However, the other two populations occurred at sites with unexpectedly low conspecific but high fathead minnow [Pimephales promelas (Rafinesque, 1820)] abundances. Further, no typical morphs were observed in either of these later two populations, contrasting with earlier research suggesting cannibalistic morphs only occur at low frequencies in salamander populations. Another anomaly of all four populations was the occurrence of cannibalistic morphs in permanent water sites, suggesting their presence was due to factors other than faster growth allowing them to occupy ephemeral habitats. Therefore, our findings suggest environmental factors inducing the cannibalistic morphism may be more complex than previously thought.
Hanifin, Charles T; Gilly, William F
Understanding the processes that generate novel adaptive phenotypes is central to evolutionary biology. We used comparative analyses to reveal the history of tetrodotoxin (TTX) resistance in TTX-bearing salamanders. Resistance to TTX is a critical component of the ability to use TTX defensively but the origin of the TTX-bearing phenotype is unclear. Skeletal muscle of TTX-bearing salamanders (modern newts, family: Salamandridae) is unaffected by TTX at doses far in excess of those that block action potentials in muscle and nerve of other vertebrates. Skeletal muscle of non-TTX-bearing salamandrids is also resistant to TTX but at lower levels. Skeletal muscle TTX resistance in the Salamandridae results from the expression of TTX-resistant variants of the voltage-gated sodium channel NaV 1.4 (SCN4a). We identified four substitutions in the coding region of salSCN4a that are likely responsible for the TTX resistance measured in TTX-bearing salamanders and variation at one of these sites likely explains variation in TTX resistance among other lineages. Our results suggest that exaptation has played a role in the evolution of the TTX-bearing phenotype and provide empirical evidence that complex physiological adaptations can arise through the accumulation of beneficial mutations in the coding region of conserved proteins.
Brockes, Jeremy P; Gates, Phillip B
Limb regeneration in adult salamanders proceeds by formation of a mound of progenitor cells called the limb blastema. It provides several pointers for regenerative medicine. These include the role of differentiated cells in the origin of the blastema, the role of regenerating axons of peripheral nerves and the importance of cell specification in conferring morphogenetic autonomy on the blastema. One aspect of regeneration that has received less attention is the ability to undergo multiple episodes without detectable change in the outcome, and with minimal effect of aging. We suggest that, although such pointers are valuable, it is important to understand why salamanders are the only adult tetrapod vertebrates able to regenerate their limbs. Although this remains a controversial issue, the existence of salamander-specific genes that play a significant role in the mechanism of regeneration provides evidence for the importance of local evolution, rather than a purely ancestral mechanism. The three-finger protein called Prod1 is discussed in the present article as an exemplar of this approach.
Pierson, Todd W.; Mckee, Anna; Spear, Stephen F.; Maerz, John C.; Camp, Carlos D.; Glenn, Travis C.
The isolation and identification of environmental DNA (eDNA) offers a non-invasive and efficient method for the detection of rare and secretive aquatic wildlife, and it is being widely integrated into inventory and monitoring efforts. The Patch-Nosed Salamander (Urspelerpes brucei) is a tiny, recently discovered species of plethodontid salamander known only from headwater streams in a small region of Georgia and South Carolina. Here, we present results of a quantitative PCR-based eDNA assay capable of detecting Urspelerpes in more than 75% of 33 samples from five confirmed streams. We deployed the method at 31 additional streams and located three previously undocumented populations of Urspelerpes. We compare the results of our eDNA assay with our attempt to use aquatic leaf litterbags for the rapid detection of Urspelerpes and demonstrate the relative efficacy of the eDNA assay. We suggest that eDNA offers great potential for use in detecting other aquatic and semi-aquatic plethodontid salamanders.
Pough, F.H.; Wilson, R.E.
The two species of mole salamander that occur in the Ithaca, New York, region (Ambystoma maculatum and A. jeffersonianum) breed in temporary ponds that are formed by accumulation of melted snow and spring rains. Water in many of these pools during the breeding season is acid; pH values as low as 3.5 have been measured. In laboratory experiments A. maculatum tolerated pHs from 6 to 10 and had greatest hatching success at pH 7 to 9. Ambystoma jeffersonianum tolerated pH 4 to 8 and was most successful at pH 5 to 6. Mortality rose abruptly beyond the tolerance limits. The pH optimum shifted upward with increasing temperature for A. jeffersonianum and downward for A. maculatum. Judging from our laboratory studies, the acidity measured in breeding ponds should cause mortality in A. maculatum and permit normal development in A. jeffersonianum. In a 4 yr study of a large, acidic vernal pond, 938 adult A. maculatum produced 486 metamorphosed juveniles (0.52 juvenile/adult), while 686 adult A. jeffersonianum produced 2157 juveniles (3.14 juveniles/adult). Because the effects of acid precipitation on the salamanders' breeding ponds are cumulative from year to year, profound changes in the salamander populations can be anticipated.
Pough, R.H.; Wilson, R.E.
The two species of mole salamander that occur in the Ithaca, New York, region (Ambystoma maculatum and A. jeffersonianum) breed in temporary ponds that are formed by accumulation of melted snow and spring rains. Water in many of these pools during the breeding season is acid; pH values as low as 3.5 have been measured. In laboratory experiments A. maculatum tolerated pHs from 6 to 10 and had greatest hatching success at pH 7 to 9. Ambystoma Jeffersonianum tolerated pH 4 to 8 and was most successful at pH 5 to 6. Mortality rose abruptly beyond the tolerance limits. The pH optimum shifted upward with increasing temperature for A. jeffersonianum and downward for A. maculatum. Judging from our laboratory studies, the acidity measured in breeding ponds should cause mortality in A. maculatum and permit normal development in A. jeffersonianum. In a four-year study of a large acidic vernal pond, 938 adult A. maculatum produced 486 metamorphosed juveniles (0.52 juvenile/adult), while 686 adult A. jeffersonianum produced 2157 juveniles (3.14 juveniles/adult). Because the effects of acid precipitation on the salamanders' breeding ponds are cumulative from year to year, profound changes in the salamander populations can be anticipated.
Hannah L. Clipp
Full Text Available Salamanders and riparian forests are intimately interconnected. Salamanders are integral to ecosystem functions, contributing to vertebrate biomass and complex food webs in riparian forests. In turn, these forests are critical ecosystems that perform many environmental services, facilitate high biodiversity and species richness, and provide habitat to salamander populations. Due to the global decline of amphibians, it is important to understand, as thoroughly and holistically as possible, the roles of environmental parameters and the impact of human activities on salamander abundance and diversity in riparian forests. To determine the population responses of salamanders to a variety of environmental factors and anthropogenic activities, we conducted a review of published literature that compared salamander abundance and diversity, and then summarized and synthesized the data into general patterns. We identify stream quality, leaf litter and woody debris, riparian buffer width, and soil characteristics as major environmental factors influencing salamander populations in riparian forests, describe and explain salamander responses to those factors, and discuss the effects of anthropogenic activities such as timber harvest, prescribed fires, urbanization, road construction, and habitat fragmentation. This review can assist land and natural resource managers in anticipating the consequences of human activities and preparing strategic conservation plans.
Burke, John N.; Bergeron, Christine M.; Todd, Brian D. [Wildlife Ecotoxicology and Physiological Ecology Program, Department of Fisheries and Wildlife Sciences, Virginia Polytechnic Institute and State University, 100 Cheatham Hall, Blacksburg, VA 24061 (United States); Hopkins, William A., E-mail: email@example.com [Wildlife Ecotoxicology and Physiological Ecology Program, Department of Fisheries and Wildlife Sciences, Virginia Polytechnic Institute and State University, 100 Cheatham Hall, Blacksburg, VA 24061 (United States)
Mercury (Hg) causes a range of deleterious effects in wildlife, but little is known about its effects on amphibians. Our objective was to determine whether Hg affects performance and behavior in two-lined salamanders (Eurycea bislineata). We collected salamanders from Hg-contaminated and reference sites and assessed speed, responsiveness, and prey capture ability. Mercury concentrations were >17x higher in salamanders from the contaminated sites and were among the highest documented in amphibians. In the first, but not in the second, locomotion trial, we found a significant effect of Hg on speed and responsiveness. In the prey capture experiment, reference salamanders ate approximately twice as many prey items as the contaminated salamanders. Together, our results suggest that sublethal Hg concentrations may negatively affect salamanders by reducing their ability to successfully execute tasks critical to survival. Future work is warranted to determine whether Hg has other sublethal effects on salamanders and whether other amphibians are similarly affected. - Mercury contamination may alter behavior and performance in the northern two-lined salamander (Eurycea bislineata).
Maliarchuk, B A
Sequence analysis of the cytochrome b gene fragment in the salamanders of the genus Salamandrella, Siberian salamander and Schrenk salamander was performed with the purpose to elucidate the effect of natural selection on the evolution of mitochondrial DNA (mtDNA) in these species. It was demonstrated that despite of notable influence of negative selection (expressed as very low dN/dS values), speciation and intraspecific divergence in salamanders was accompanied by the appearance of radical amino acid substitutions, caused by the influence of positive (directional) selection. To examine the evolutionary pattern of synonymous mtDNA sites, distribution of conservative and non-conservative substitutions was analyzed. The rates of conservative and non-conservative substitutions were nearly equal, pointing to neutrality of mutation process at synonymous mtDNA sites of salamanders. Analysis of conservative and non-conservative synonymous substitution distributions in different parts of phylogenetic trees showed that the differences between the synonymous groups compared were statistically significant only in one phylogenetic group of Siberian salamander (haplogroup C) (P = 0.02). In the group of single substitutions, located at terminal phylogenetic branches of Siberian salamanders from group C, increased rate of conservative substitutions was observed. Based on these findings, it was suggested that selective processes could have an influence on the formation of the synonymous substitution profile in the Siberian salamander mtDNA fragment examined.
Eric J. Gustafson; Nathan L. Murphy; Thomas R. Crow
A GIS model predicting the spatial distribution of terrestrial salamander abundance based on topography and forest age was developed using parameters derived from the literature. The model was tested by sampling salamander abundance across the full range of site conditions used in the model. A regression of the predictions of our GIS model against these sample data...
F. Harvey Pough; Donald H. Rhodes; Andres Collazo
Because of the importance of salamanders in forest food chains, the effects of forest management practices on populations of these animals warrant consideration. We compared the numbers and activity patterns of salamanders in areas of a deciduous forest in central New York State that had been cut selectively for firewood, or c1earcut, or planted with conifers. Numbers...
Link, William; Hesed, Kyle Miller
Knowledge of organisms’ growth rates and ages at sexual maturity is important for conservation efforts and a wide variety of studies in ecology and evolutionary biology. However, these life history parameters may be difficult to obtain from natural populations: individuals encountered may be of unknown age, information on age at sexual maturity may be uncertain and interval-censored, and growth data may include both individual heterogeneity and measurement errors. We analyzed mark–recapture data for Red-backed Salamanders (Plethodon cinereus) to compare sex-specific growth rates and ages at sexual maturity. Aging of individuals was made possible by the use of a von Bertalanffy model of growth, complemented with models for interval-censored and imperfect observations at sexual maturation. Individual heterogeneity in growth was modeled through the use of Gamma processes. Our analysis indicates that female P. cinereus mature earlier and grow more quickly than males, growing to nearly identical asymptotic size distributions as males.
Diekwisch, Thomas G H; Jin, Tianquan; Wang, Xinping; Ito, Yoshihiro; Schmidt, Marcella; Druzinsky, Robert; Yamane, Akira; Luan, Xianghong
Amelogenins are the major proteins involved in tooth enamel formation. In the present study, we have cloned and sequenced four novel amelogenins from three amphibian species in order to analyze similarities and differences between mammalian and non-mammalian amelogenins. The newly sequenced amphibian amelogenin sequences were from a red-eyed tree frog (Litoria chloris) and a Mexican axolotl (Ambystoma mexicanum). We identified two amelogenin isoforms in the Eastern red-backed salamander (Plethodon cinereus). Sequence comparisons confirmed that non-mammalian amelogenins are overall shorter than their mammalian counterparts, contain less proline and less glutamine, and feature shorter polyproline tripeptide repeat stretches than mammalian amelogenins. We propose that unique sequence parameters of mammalian amelogenins might be a pre-requisite for complex mammalian enamel prism architecture. Copyright (c) 2009 S. Karger AG, Basel.
Jon M. Davenport
Full Text Available Recent research indicates that social and environmental factors influence egg and clutch sizes in amphibians. However, most of this work is based on the reproductively diverse order Anura (frogs and toads, whereas less research has been conducted on Caudata (salamanders and Gymnophiona (caecilians. Researchers have suggested that a relationship exists between social and environmental factors and egg and clutch sizes in salamanders, but studies controlling for phylogenetic context are lacking. We could not identify a sufficient number of comparisons for social influences on egg and clutch sizes; therefore, we focused on environmental influences for this study. Data on egg size, clutch size, environmental factors, and phylogenies for salamanders were assembled from the scientific literature. We used independent, pair-wise comparisons to investigate the association of larval salamander habitat and egg size and the association of larval salamander habitat with clutch sizes within a phylogenetic framework. There is a significant association between larval habitat and egg size; specifically, stream-breeding species produce larger eggs. There is no significant association between larval habitat and clutchsize. Our study confirms earlier reports that salamander egg size is associated with larval environments, but is the first to use phylogenetically independent contrasts to account for the lack of phylogenetic independence of the traits measured (egg size and clutch size associated with many of the diverse lineages. Our study shows that environmental selection pressure can be quite strong on one aspect of salamander reproduction—egg size.
Musgrave, Maria A. [Los Alamos National Lab. (LANL), Los Alamos, NM (United States). Wildlife Management
This is an article from the Los Alamos Living magazine. Los Alamos National Laboratory sits on a beautiful and unique landscape that provides important protected habitat to many species, including a few that are federally-listed as threatened or endangered. These species are the Jemez Mountains Salamander, the Mexican Spotted Owl, the Southwestern Willow Flycatcher, the Yellow-billed Cuckoo, and the New Mexico Meadow Jumping Mouse. Part of the job of the Laboratory's wildlife biologists is to survey for these species each year and determine what actions need to be taken if they are found.
Cabelguen, Jean-Marie; Ijspeert, Auke; Lamarque, Stéphanie; Ryczko, Dimitri
Much of what we know about the flexibility of the locomotor networks in vertebrates is derived from studies examining the adaptation of limb movements during stepping in various conditions. However, the body movements play important roles during locomotion: they produce the thrust during undulatory locomotion and they help to increase the stride length during legged locomotion. In this chapter, we review our current knowledge about the flexibility in the neuronal circuits controlling the body musculature during locomotion. We focus especially on salamander because, as an amphibian, this animal is able to display a rich repertoire of aquatic and terrestrial locomotor modes.
Full Text Available Japanese giant salamander Andrias japonicus Chordata/Vertebrata/Amphibia Andrias_japonicus_L.png Andrias_jap...onicus_NL.png Andrias_japonicus_S.png Andrias_japonicus_NS.png http://biosciencedbc....jp/taxonomy_icon/icon.cgi?i=Andrias+japonicus&t=L http://biosciencedbc.jp/taxonomy_icon/icon.cgi?i=Andrias+japon...icus&t=NL http://biosciencedbc.jp/taxonomy_icon/icon.cgi?i=Andrias+japonicus...&t=S http://biosciencedbc.jp/taxonomy_icon/icon.cgi?i=Andrias+japonicus&t=NS ...
Pough, F H
Spotted salamanders breed in temporary pools formed in early spring by melted snow and rain. Many of these pools reflect the low pH of precipitation in the northeastern United States. Egg mortality is low (less than 1 percent) in pools near neutrality, but high (greater than 60 percent) in pools more acid than pH 6. Developmental anomalies and the embryonic stage at which death occurs are the same in field situations as at corresponding pH's in laboratory experiments.
Jessica A. Homyack
Full Text Available Intensively-managed forest (IMF ecosystems support environmental processes, retain biodiversity and reduce pressure to extract wood products from other forests, but may affect species, such as plethodontid salamanders, that are associated with closed canopies and possess limited vagility. We describe: (1 critical aspects of IMF ecosystems; (2 effectiveness of plethodontid salamanders as barometers of forest change; (3 two case studies of relationships between salamanders and coarse woody debris (CWD; and (4 research needs for effective management of salamanders in IMF ecosystems. Although plethodontid salamanders are sensitive to microclimate changes, their role as ecological indicators rarely have been evaluated quantitatively. Our case studies of CWD and salamanders in western and eastern forests demonstrated effects of species, region and spatial scale on the existence and strength of relationships between plethodontid species and a “critical” microhabitat variable. Oregon slender salamanders (Batrachoseps wrighti were more strongly associated with abundance of CWD in managed second growth forests than ensatina salamanders (Ensatina eschscholtzii. Similarly, CWD was not an important predictor of abundance of Appalachian salamanders in managed hardwood forest. Gaining knowledge of salamanders in IMF ecosystems is critical to reconciling ecological and economic objectives of intensive forest management, but faces challenges in design and implementation.
Sessions, Stanley K; Bizjak Mali, Lilijana; Green, David M; Trifonov, Vladimir; Ferguson-Smith, Malcolm
A major goal of genomic and reproductive biology is to understand the evolution of sex determination and sex chromosomes. Species of the 2 genera of the Salamander family Proteidae - Necturus of eastern North America, and Proteus of Southern Europe - have similar-looking karyotypes with the same chromosome number (2n = 38), which differentiates them from all other salamanders. However, Necturus possesses strongly heteromorphic X and Y sex chromosomes that Proteus lacks. Since the heteromorphic sex chromosomes of Necturus were detectable only with C-banding, we hypothesized that we could use C-banding to find sex chromosomes in Proteus. We examined mitotic material from colchicine-treated intestinal epithelium, and meiotic material from testes in specimens of Proteus, representing 3 genetically distinct populations in Slovenia. We compared these results with those from Necturus. We performed FISH to visualize telomeric sequences in meiotic bivalents. Our results provide evidence that Proteus represents an example of sex chromosome turnover in which a Necturus-like Y-chromosome has become permanently translocated to another chromosome converting heteromorphic sex chromosomes to homomorphic sex chromosomes. These results may be key to understanding some unusual aspects of demographics and reproductive biology of Proteus, and are discussed in the context of models of the evolution of sex chromosomes in amphibians.
Deitloff, Jennifer; Alcorn, Michael A; Graham, Sean P
Two of the most common mating tactics in vertebrates are mate guarding and territoriality, yet much of the research on these strategies has focused on mating systems in birds, despite novel insights gained from studying less traditional systems. North American stream salamanders that comprise the Eurycea bislineata complex represent an excellent nontraditional system for comparing mating strategies because these species exhibit a continuum of male morphologies, diverse habitat associations, and various potential mating strategies. We studied two species within this complex that exhibit the extremes of this continuum, Eurycea aquatica (robust morph) and Eurycea cirrigera (slender morph). The larger head in males of E. aquatica is due to larger musculature around the jaw and may be associated with aggressive behavior. Therefore, we hypothesized that the robust morphology exhibited by males of E. aquatica provides benefits during either territorial defense or mate defense and that males of E. cirrigera would not exhibit aggression in either scenario. We found that neither species exhibited aggressive behavior to defend a territory. However, in the presence of a female, males of E. aquatica were significantly more aggressive toward intruding males than were males of E. cirrigera. Therefore, mate-guarding behavior occurs in E. aquatica, and the enlarged head of males likely aids in deterring rivals. This is the first demonstration of mate-guarding behavior in a plethodontid, the most speciose family of salamanders.
Karraker, Nancy E., E-mail: firstname.lastname@example.org [Department of Environmental and Forest Biology, State University of New York, College of Environmental Science and Forestry, Syracuse, NY 13210 (United States); Gibbs, James P. [Department of Environmental and Forest Biology, State University of New York, College of Environmental Science and Forestry, Syracuse, NY 13210 (United States)
It has been postulated that road deicing salts are sufficiently diluted by spring rains to ameliorate any physiological impacts to amphibians breeding in wetlands near roads. We tested this conjecture by exposing clutches of the spotted salamander (Ambystoma maculatum) to three chloride concentrations (1 mg/L, 145 mg/L, 945 mg/L) for nine days, then transferred clutches to control water for nine days, and measured change in mass at three-day intervals. We measured mass change because water uptake by clutches reduces risks to embryos associated with freezing, predation, and disease. Clutches in controls sequestered water asymptotically. Those in the moderate concentrations lost 18% mass initially and regained 14% after transfer to control water. Clutches in high concentration lost 33% mass and then lost an additional 8% after transfer. Our results suggest that spring rains do not ameliorate the effects of deicing salts in wetlands with extremely high chloride concentrations. - Road deicing salts irreversibly disrupts osmoregulation of salamander egg clutches.
Godwin, James W; Pinto, Alexander R; Rosenthal, Nadia A
The failure to replace damaged body parts in adult mammals results from a muted growth response and fibrotic scarring. Although infiltrating immune cells play a major role in determining the variable outcome of mammalian wound repair, little is known about the modulation of immune cell signaling in efficiently regenerating species such as the salamander, which can regrow complete body structures as adults. Here we present a comprehensive analysis of immune signaling during limb regeneration in axolotl, an aquatic salamander, and reveal a temporally defined requirement for macrophage infiltration in the regenerative process. Although many features of mammalian cytokine/chemokine signaling are retained in the axolotl, they are more dynamically deployed, with simultaneous induction of inflammatory and anti-inflammatory markers within the first 24 h after limb amputation. Systemic macrophage depletion during this period resulted in wound closure but permanent failure of limb regeneration, associated with extensive fibrosis and disregulation of extracellular matrix component gene expression. Full limb regenerative capacity of failed stumps was restored by reamputation once endogenous macrophage populations had been replenished. Promotion of a regeneration-permissive environment by identification of macrophage-derived therapeutic molecules may therefore aid in the regeneration of damaged body parts in adult mammals.
Zhang, Hui; Yu, Pengcheng; Zhong, Shengwei; Ge, Tingting; Peng, Shasha; Guo, Xiaoquan; Zhou, Zuohong
Telocytes (TCs), novel interstitial cells, have been identified in various organs of many mammals. However, information about TCs of lower animals remains rare. Herein, pancreatic TCs of the Chinese giant salamanders (Andrias davidianus) were identified by CD34 immunohistochemistry (IHC) and transmission electron microscopy (TEM). The IHC micrographs revealed CD34(+) TCs with long telopodes (Tps) that were located in the interstitium of the pancreas. CD34(+) TCs/Tps were frequently observed between exocrine acinar cells and were close to blood vessels. The TEM micrographs also showed the existence of TCs in the interstitium of the pancreas. TCs had distinctive ultrastructural features, such as one to three very long and thin Tps with podoms and podomers, caveolae, dichotomous branching, neighbouring exosomes and vesicles. The Tps and exosomes were found in close proximity to exocrine acinar cells and α cells. It is suggested that TCs may play a role in the regeneration of acinar cells and α cells. In conclusion, our results demonstrated the presence of TCs in the pancreas of the Chinese giant salamander. This finding will assist us in a better understanding of TCs functions in the amphibian pancreas. © 2016 The Authors. Journal of Cellular and Molecular Medicine published by John Wiley & Sons Ltd and Foundation for Cellular and Molecular Medicine.
We investigated the metapopulation structure of the California tiger salamander (Ambystoma californiense) using a combination of indirect and direct methods to evaluate two key requirements of modern metapopulation models: 1) that patches support somewhat independent populations ...
US Fish and Wildlife Service, Department of the Interior — This is a report that outlines the results of a one-day survey for the endangered Cheat Mountain Salamander at Canaan Valley National Wildlife Refuge in 2004. The...
Regulatory agencies need rapid indicators of hydrologic permanence for jurisdictional determinations of headwater streams. Our study objective was to assess the utility of larval salamander presence and assemblage structure and habitat variables for determining stream permanence ...
US Fish and Wildlife Service, Department of the Interior — Aquatic habitat of the endangered Barton Springs salamander, Eurycea sosorum, in Travis County, Texas can potentially be impacted by contaminants in surface runoff...
US Fish and Wildlife Service, Department of the Interior — From 2000 to 2008 data was collected from three Cheat Mountain salamander coverboard sites at Canaan Valley National Wildlife Refuge: Cabin Knob A, Cabin Knob B, and...
US Fish and Wildlife Service, Department of the Interior — This documents six different surveys between July and September, 2008 that were done to monitor the endangered Cheat Mountain salamander at Canaan Valley National...
U.S. Environmental Protection Agency — Stoichiometry of excreta and excretion rates of a stream-dwelling plethodontid salamander in Cincinnati, OH, USA. This dataset is associated with the following...
Kändler, Tiit, 1948-
Salamander on selgroogsete seas ainulaadne olevus. Ta suudab endale kasvatada ka täiskasvanuna uued kehaosad. Kui arstiteadlased välja uurivad, kuidas ta seda teeb, võib see aidata ka inimest rasketest haavadest ravida
Among vertebrates, urodele amphibians are the only tetrapods with the ability to regenerate complex structures such as limbs, tail, and spinal cord throughout their lives. Furthermore, the salamander regeneration process has been shown to reverse tumorigenicity. Fibroblasts are essential for salamander regeneration, but the mechanisms underlying their role in the formation of a regeneration blastema remain unclear. Here, I review the role of fibroblasts in salamander limb regeneration and how their activity compares with that of human fibroblasts. In addition, the question of whether salamander blastema tissue could induce regeneration and tumor regression in animals with a limited regeneration ability is discussed. A deeper understanding of these processes may lead to the development of novel regenerative and anticancer therapies.
US Fish and Wildlife Service, Department of the Interior — Many springs associated with the Edwards Aquifer of Texas are inhabited by relict populations of neotenic salamanders in the genus Eurycea. This study was done to...
Full Text Available We report new aquatic predators feeding on Northern spectacled salamander eggs, Salamandrina perspicillata, an endemic Italian species. Eggs were preyed upon by the leech, Trocheta bykowskii, and the trichopteran larvae of Potamophylax cingulatus and Halesus appenninus.
subflavus Red bat Lasiurus borealis Hoary bat Lasiurus cinereus Evening bat Nycticeius humeralis Big-eared bat Plectus reainesquii Free-tailed bat Tadarida...salamander Ambystoma tigrinum Dusky salamander Desmognathus fuscus M Many-lined salamander Stereochilus marginatus Slimy salamander Plethodon glutinosus
Kundey, Shannon M A; Millar, Roberto; McPherson, Justin; Gonzalez, Maya; Fitz, Aleyna; Allen, Chadbourne
We explored tiger salamanders' (Ambystoma tigrinum) learning to execute a response within a maze as proximal visual cue conditions varied. In Experiment 1, salamanders learned to turn consistently in a T-maze for reinforcement before the maze was rotated. All learned the initial task and executed the trained turn during test, suggesting that they learned to demonstrate the reinforced response during training and continued to perform it during test. In a second experiment utilizing a similar procedure, two visual cues were placed consistently at the maze junction. Salamanders were reinforced for turning towards one cue. Cue placement was reversed during test. All learned the initial task, but executed the trained turn rather than turning towards the visual cue during test, evidencing response learning. In Experiment 3, we investigated whether a compound visual cue could control salamanders' behaviour when it was the only cue predictive of reinforcement in a cross-maze by varying start position and cue placement. All learned to turn in the direction indicated by the compound visual cue, indicating that visual cues can come to control their behaviour. Following training, testing revealed that salamanders attended to stimuli foreground over background features. Overall, these results suggest that salamanders learn to execute responses over learning to use visual cues but can use visual cues if required. Our success with this paradigm offers the potential in future studies to explore salamanders' cognition further, as well as to shed light on how features of the tiger salamanders' life history (e.g. hibernation and metamorphosis) impact cognition.
Full Text Available Captive rearing and reintroduction / translocation are increasingly used as tools to supplement wild populations of threatened species. Reintroducing captive-reared Chinese giant salamanders may help to augment the declining wild populations and conserve this critically endangered amphibian. We released 31 captive-reared juvenile giant salamanders implanted with VHF radio transmitters at the Heihe River (n = 15 and the Donghe River (n = 16 in the Qinling Mountains of central China. Salamanders were monitored every day for survival from April 28th 2013 to September 3rd 2014. We attempted to recapture all living individuals by the end of the study, measured their body mass and total body length, and checked for abnormalities and presence of external parasites. Two salamanders at the Heihe River and 10 animals at the Donghe River survived through the project timeline. Nine salamanders were confirmed dead, while the status of the other 10 animals was undetermined. The annual survival rate of giant salamanders at the Donghe River (0.702 was 1.7-fold higher than that at the Heihe River (0.405. Survival increased as individuals were held longer following surgery, whereas body mass did not have a significant impact on survival rate. All salamanders recaptured from the Donghe River (n = 8 increased in mass (0.50 ± 0.13 kg and length (5.5 ± 1.5 cm after approximately 11 months in the wild, and they were only 7% lighter than wild animals of the same length (mean residual = -0.033 ± 0.025. Our results indicate that captive-reared Chinese giant salamanders can survive in the wild one year after release and adequate surgical recovery time is extremely important to post-release survival. Future projects may reintroduce older juveniles to achieve better survival and longer monitoring duration.
Zhang, Lu; Jiang, Wei; Wang, Qi-Jun; Zhao, Hu; Zhang, Hong-Xing; Marcec, Ruth M.; Willard, Scott T.; Kouba, Andrew J.
Captive rearing and reintroduction / translocation are increasingly used as tools to supplement wild populations of threatened species. Reintroducing captive-reared Chinese giant salamanders may help to augment the declining wild populations and conserve this critically endangered amphibian. We released 31 captive-reared juvenile giant salamanders implanted with VHF radio transmitters at the Heihe River (n = 15) and the Donghe River (n = 16) in the Qinling Mountains of central China. Salamanders were monitored every day for survival from April 28th 2013 to September 3rd 2014. We attempted to recapture all living individuals by the end of the study, measured their body mass and total body length, and checked for abnormalities and presence of external parasites. Two salamanders at the Heihe River and 10 animals at the Donghe River survived through the project timeline. Nine salamanders were confirmed dead, while the status of the other 10 animals was undetermined. The annual survival rate of giant salamanders at the Donghe River (0.702) was 1.7-fold higher than that at the Heihe River (0.405). Survival increased as individuals were held longer following surgery, whereas body mass did not have a significant impact on survival rate. All salamanders recaptured from the Donghe River (n = 8) increased in mass (0.50 ± 0.13 kg) and length (5.5 ± 1.5 cm) after approximately 11 months in the wild, and they were only 7% lighter than wild animals of the same length (mean residual = -0.033 ± 0.025). Our results indicate that captive-reared Chinese giant salamanders can survive in the wild one year after release and adequate surgical recovery time is extremely important to post-release survival. Future projects may reintroduce older juveniles to achieve better survival and longer monitoring duration. PMID:27258650
Fan, Yuding; Chang, Ming Xian; Ma, Jie; LaPatra, Scott E.; Hu, Yi Wei; Huang, Lili; Nie, Pin; Zeng, Lingbing
International audience; AbstractThe emergence of an infectious viral disease caused by the Chinese giant salamander iridovirus (GSIV) has led to substantial economic losses. However, no more molecular information is available for the understanding of the mechanisms associated with virus–host interaction. In this study, de novo sequencing was used to obtain abundant high-quality ESTs and investigate differentially-expressed genes in the spleen of Chinese giant salamanders that were either infe...
Zhang, Lu; Jiang, Wei; Wang, Qi-Jun; Zhao, Hu; Zhang, Hong-Xing; Marcec, Ruth M; Willard, Scott T; Kouba, Andrew J
Captive rearing and reintroduction / translocation are increasingly used as tools to supplement wild populations of threatened species. Reintroducing captive-reared Chinese giant salamanders may help to augment the declining wild populations and conserve this critically endangered amphibian. We released 31 captive-reared juvenile giant salamanders implanted with VHF radio transmitters at the Heihe River (n = 15) and the Donghe River (n = 16) in the Qinling Mountains of central China. Salamanders were monitored every day for survival from April 28th 2013 to September 3rd 2014. We attempted to recapture all living individuals by the end of the study, measured their body mass and total body length, and checked for abnormalities and presence of external parasites. Two salamanders at the Heihe River and 10 animals at the Donghe River survived through the project timeline. Nine salamanders were confirmed dead, while the status of the other 10 animals was undetermined. The annual survival rate of giant salamanders at the Donghe River (0.702) was 1.7-fold higher than that at the Heihe River (0.405). Survival increased as individuals were held longer following surgery, whereas body mass did not have a significant impact on survival rate. All salamanders recaptured from the Donghe River (n = 8) increased in mass (0.50 ± 0.13 kg) and length (5.5 ± 1.5 cm) after approximately 11 months in the wild, and they were only 7% lighter than wild animals of the same length (mean residual = -0.033 ± 0.025). Our results indicate that captive-reared Chinese giant salamanders can survive in the wild one year after release and adequate surgical recovery time is extremely important to post-release survival. Future projects may reintroduce older juveniles to achieve better survival and longer monitoring duration.
Jessica A. Homyack; Andrew J. Kroll
Intensively-managed forest (IMF) ecosystems support environmental processes, retain biodiversity and reduce pressure to extract wood products from other forests, but may affect species, such as plethodontid salamanders, that are associated with closed canopies and possess limited vagility. We describe: (1) critical aspects of IMF ecosystems; (2) effectiveness of plethodontid salamanders as barometers of forest change; (3) two case studies of relationships between salamanders and coarse wood...
Lunghi, Enrico; Manenti, Raoul; Canciani, Giancarlo; Scarì, Giorgio; Pennati, Roberta; Ficetola, Gentile Francesco
Information on species thermal physiology is extremely important to understand species responses to environmental heterogeneity and changes. Thermography is an emerging technology that allows high resolution and accurate measurement of body temperature, but until now it has not been used to study thermal physiology of amphibians in the wild. Hydromantes terrestrial salamanders are strongly depending on ambient temperature for their activity and gas exchanges, but information on their body temperature is extremely limited. In this study we tested if Hydromantes salamanders are thermoconform, we assessed whether there are temperature differences among body regions, and evaluated the time required to reach the thermal equilibrium. During summers of 2014 and 2015 we analysed 56 salamanders (Hydromantes ambrosii and Hydromantes italicus) using infrared thermocamera. We photographed salamanders at the moment in which we found them and 1, 2, 3, 4, 5 and 15min after having kept them in the hands. Body temperature was equal to air temperature; salamanders attained the equilibrium with air temperature in about 8min, the time required to reach equilibrium was longer in individuals with large body size. We detected small temperature differences between body parts, the head being slightly warmer than the body and the tail (mean difference: 0.05°C). These salamanders quickly reach the equilibrium with the environment, thus microhabitat measurement allows obtaining accurate information on their tolerance limits.
Chevallier, Stéphanie; Jan Ijspeert, Auke; Ryczko, Dimitri; Nagy, Frédéric; Cabelguen, Jean-Marie
Among living tetrapods, salamanders are regarded as most closely resembling the first terrestrial vertebrates, and are therefore an interesting group in which the evolutionary changes in the locomotor behaviour from aquatic to terrestrial habitats can be inferred. Salamanders exhibit two locomotor modes: swimming and terrestrial stepping. The swimming is anguilliform and resembles closely that of the lamprey. On the ground, the salamander switches to a stepping gait with axial undulations that is also observed in many reptiles. The salamander is therefore ideally suited for examining the neural mechanisms for the generation of these two locomotor modes, as well as the neural mechanisms of gait transition. In the present paper, we describe the kinematics and patterns of activation of axial and limb muscles during stepping and swimming in adult salamanders. We then review the current neurobiological data about the organisation of the spinal networks underlying swimming and stepping, and the mechanisms of gait transition. Finally we report modelling studies aimed at understanding the organisation and operation of the salamander locomotor circuits. Altogether, the neurobiological and the modelling data support the hypothesis of a phylogenetic conservatism from agnathians to amphibians of the spinal locomotor networks generating axial motor patterns.
Karraker, Nancy E; Gibbs, James P
It has been postulated that road deicing salts are sufficiently diluted by spring rains to ameliorate any physiological impacts to amphibians breeding in wetlands near roads. We tested this conjecture by exposing clutches of the spotted salamander (Ambystoma maculatum) to three chloride concentrations (1 mg/L, 145 mg/L, 945 mg/L) for nine days, then transferred clutches to control water for nine days, and measured change in mass at three-day intervals. We measured mass change because water uptake by clutches reduces risks to embryos associated with freezing, predation, and disease. Clutches in controls sequestered water asymptotically. Those in the moderate concentrations lost 18% mass initially and regained 14% after transfer to control water. Clutches in high concentration lost 33% mass and then lost an additional 8% after transfer. Our results suggest that spring rains do not ameliorate the effects of deicing salts in wetlands with extremely high chloride concentrations.
Kumar, Anoop; Gates, Phillip B; Brockes, Jeremy P
Limb regeneration in larval and adult salamanders proceeds from a mound of mesenchymal stem cells called the limb blastema. The blastema gives rise just to those structures distal to its level of origin, and this property of positional identity is reset to more proximal values by treatment with retinoic acid. We have identified a cell surface protein, called Prod1/CD59, which appears to be a determinant of proximodistal identity. Prod1 is expressed in an exponential gradient in an adult limb as determined by detection of both mRNA and immunoreactive protein. Prod1 protein is up-regulated after treatment of distal blastemas with RA and this is particularly marked in cells of the dermis. These cells have previously been implicated in pattern formation during limb regeneration.
T.M. Luhring; C.A. Young
Although salamanders are excellent indicators of environmental health, the ability to catch them efficiently without substantially disrupting their habitat is not always practical or even possible with current techniques. Ripping open logs and raking leaf packs onto shore (Bruce 1972) are examples of such practices that are disruptive but widely used by herpetologists who have no other means of efficient collection. Drift fences with pitfall traps are effective in catching animals moving within or between habitats but are time consuming and require an initial financial investment and constant upkeep to maintain functionality and prevent animal fatalities (Gibbons and Semlitsch 1981). One current alternative to drift fences is the use of coverboards (Grant et al. 1992), which require less maintenance and sampling effort than drift fences. However, coverboards do not integrate captures over a long time period and often result in a lower number of captures per trap (Grant et al. 1992).
Caldwell, J P; Thorp, J H; Jervey, T O
Tadpoles of the barking tree frog, Hyla gratiosa, are abundant in spring and summer in some ponds and Carolina bays on the Savannah River Plant near Aiken, South Carolina. To determine how these tadpoles survive in the presence of predaceous salamander larvae, Ambystoma talpoideum, and larvae of an aeshnid dragonfly, Anax junius, we determined fields densities and sizes of the predators and the prey and conducted predation experiments in the laboratory. Tadpoles rapidly grow to a size not captured by Ambystoma, although Anax larvae can capture slightly larger tadpoles. Differing habitat preferences among the tadpoles and the two predator species probably aid in reducing predation pressure. Preliminary work indicates that the tadpoles may have an immobility response to an attack by a predator. In addition, the smallest, most vulnerable tadpoles have a distinctive color pattern which may function to disrupt the body outline and make them indiscernable to predators.
Becker, Matthew H.; Brucker, Robert M.; Schwantes, Christian R.; Harris, Reid N.; Minbiole, Kevin P. C.
The disease chytridiomycosis, which is caused by the chytrid fungus Batrachochytrium dendrobatidis, is associated with recent declines in amphibian populations. Susceptibility to this disease varies among amphibian populations and species, and resistance appears to be attributable in part to the presence of antifungal microbial species associated with the skin of amphibians. The betaproteobacterium Janthinobacterium lividum has been isolated from the skins of several amphibian species and produces the antifungal metabolite violacein, which inhibits B. dendrobatidis. In this study, we added J. lividum to red-backed salamanders (Plethodon cinereus) to obtain an increased range of violacein concentrations on the skin. Adding J. lividum to the skin of the salamander increased the concentration of violacein on the skin, which was strongly associated with survival after experimental exposure to B. dendrobatidis. As expected from previous work, some individuals that did not receive J. lividum and were exposed to B. dendrobatidis survived. These individuals had concentrations of bacterially produced violacein on their skins that were predicted to kill B. dendrobatidis. Our study suggests that a threshold violacein concentration of about 18 μM on a salamander's skin prevents mortality and morbidity caused by B. dendrobatidis. In addition, we show that over one-half of individuals in nature support antifungal bacteria that produce violacein, which suggests that there is a mutualism between violacein-producing bacteria and P. cinereus and that adding J. lividum is effective for protecting individuals that lack violacein-producing skin bacteria. PMID:19717627
Becker, Matthew H; Brucker, Robert M; Schwantes, Christian R; Harris, Reid N; Minbiole, Kevin P C
The disease chytridiomycosis, which is caused by the chytrid fungus Batrachochytrium dendrobatidis, is associated with recent declines in amphibian populations. Susceptibility to this disease varies among amphibian populations and species, and resistance appears to be attributable in part to the presence of antifungal microbial species associated with the skin of amphibians. The betaproteobacterium Janthinobacterium lividum has been isolated from the skins of several amphibian species and produces the antifungal metabolite violacein, which inhibits B. dendrobatidis. In this study, we added J. lividum to red-backed salamanders (Plethodon cinereus) to obtain an increased range of violacein concentrations on the skin. Adding J. lividum to the skin of the salamander increased the concentration of violacein on the skin, which was strongly associated with survival after experimental exposure to B. dendrobatidis. As expected from previous work, some individuals that did not receive J. lividum and were exposed to B. dendrobatidis survived. These individuals had concentrations of bacterially produced violacein on their skins that were predicted to kill B. dendrobatidis. Our study suggests that a threshold violacein concentration of about 18 microM on a salamander's skin prevents mortality and morbidity caused by B. dendrobatidis. In addition, we show that over one-half of individuals in nature support antifungal bacteria that produce violacein, which suggests that there is a mutualism between violacein-producing bacteria and P. cinereus and that adding J. lividum is effective for protecting individuals that lack violacein-producing skin bacteria.
Johnson, Mark S; McFarland, Craig A; Bazar, Matthew A; Quinn, Michael J; LaFiandra, Emily May; Talent, Larry G
The explosive, octahydro-1,3,5,7-tetranitro-1,3,5,7-tetrazocine or high-melting explosive (HMX), has been found in soils in areas used for testing and training by the military. Many of these areas contain habitat for valued wildlife species. In an effort to better understand the environmental consequences from exposure, a reptilian (western fence lizard [Sceloporus occidentalis]), an amphibian (red-backed salamander [Plethodon cinereus]), and a mammalian species (rabbit [Oryctolagus cuniculus]) were exposed to HMX under controlled laboratory conditions. Lizards and rabbits were exposed to HMX by way of corn oil through gavage, and salamanders were exposed to HMX in soil. Two deaths occurred from acute oral exposures to lizards to 5000 mg HMX/kg BW. Histological and gross pathologic assessment suggested gut impaction as a possible cause of death. Salamanders exposed to concentrations of HMX in soil 24 h after oral exposures. An LD(50) for rabbits was calculated as 93 mg/kg (95% confidence interval 76-117). A subacute 14-day testing regime found a lowest observed effect level of 10 mg/kg-d and a no observed adverse effect level of 5 mg/kg-d based on hyperkinesia and seizure incidence, although changes suggesting functional hepatic alterations were also found. These data suggest that physiologic differences between species, particularly in gastrointestinal structure and function, can affect the absorption of HMX and hence lead to marked differences in toxicity from exposure to the same compound.
The effects of the insecticides Guthion (technical grade) and Guthion 2S(commercial formulation) on survival and growth of tadpoles of the Pacific treefrog Pseudacris regilla, and larvae of the Northwestern salamander Ambystoma gracile and the spotted salamander Ambystoma macula...
Alexandrino, J.; Ferrand, N.; Arntzen, J.W.
Morphometric and colour pattern variation in the endemic Iberian salamander Chioglossa lusitanica is concordant with the genetic differentiation of two groups of populations separated by the Mondego river in Portugal. Salamanders from the south have shorter digits than those from the north. Clinal v
Grant, Evan H. Campbell; Wiewel, Amber N. M.; Rice, Karen C.
Stream ecosystems are particularly sensitive to urbanization, and tolerance of water-quality parameters is likely important to population persistence of stream salamanders. Forecasted climate and landscape changes may lead to significant changes in stream flow, chemical composition, and temperatures in coming decades. Protected areas where landscape alterations are minimized will therefore become increasingly important for salamander populations. We surveyed 29 streams at three national parks in the highly urbanized greater metropolitan area of Washington, DC. We investigated relationships among water-quality variables and occupancy of three species of stream salamanders (Desmognathus fuscus, Eurycea bislineata, and Pseudotriton ruber). With the use of a set of site-occupancy models, and accounting for imperfect detection, we found that stream-water temperature limits salamander occupancy. There was substantial uncertainty about the effects of the other water-quality variables, although both specific conductance (SC) and pH were included in competitive models. Our estimates of occupancy suggest that temperature, SC, and pH have some importance in structuring stream salamander distribution.
Kenison, Erin K.; Litt, Andrea R.; Pilliod, David; McMahon, Thomas E.
Predators can influence prey directly through consumption or indirectly through nonconsumptive effects (NCEs) by altering prey behavior, morphology, and life history. We investigated whether predator-avoidance behaviors by larval long-toed salamanders (Ambystoma macrodactylum) in lakes with nonnative trout result in NCEs on morphology and development. Field studies in lakes with and without trout were corroborated by experimental enclosures, where prey were exposed only to visual and chemical cues of predators. We found that salamanders in lakes with trout were consistently smaller than in lakes without trout: 38% lower weight, 24% shorter body length, and 29% shorter tail length. Similarly, salamanders in protective enclosures grew 2.9 times slower when exposed to visual and olfactory trout cues than when no trout cues were present. Salamanders in trout-free lakes and enclosures were 22.7 times and 1.48 times, respectively, more likely to metamorphose during the summer season than those exposed to trout in lakes and/or their cues. Observed changes in larval growth rate and development likely resulted from a facultative response to predator-avoidance behavior and demonstrate NCEs occurred even when predation risk was only perceived. Reduced body size and growth, as well as delayed metamorphosis, could have ecological consequences for salamander populations existing with fish if those effects carry-over into lower recruitment, survival, and fecundity.
Frahry, Matthew Blake; Sun, Cheng; Chong, Rebecca A; Mueller, Rachel Lockridge
Across the tree of life, species vary dramatically in nuclear genome size. Mutations that add or remove sequences from genomes-insertions or deletions, or indels-are the ultimate source of this variation. Differences in the tempo and mode of insertion and deletion across taxa have been proposed to contribute to evolutionary diversity in genome size. Among vertebrates, most of the largest genomes are found within the salamanders, an amphibian clade with genome sizes ranging from ~14 to ~120 Gb. Salamander genomes have been shown to experience slower rates of DNA loss through small (i.e., salamander genomes resulting from larger deletions. Here, we focus on one type of large deletion-ectopic-recombination-mediated removal of LTR retrotransposon sequences. In ectopic recombination, double-strand breaks are repaired using a "wrong" (i.e., ectopic, or non-allelic) template sequence-typically another locus of similar sequence. When breaks occur within the LTR portions of LTR retrotransposons, ectopic-recombination-mediated repair can produce deletions that remove the internal transposon sequence and the equivalent of one of the two LTR sequences. These deletions leave a signature in the genome-a solo LTR sequence. We compared levels of solo LTRs in the genomes of four salamander species with levels present in five vertebrates with smaller genomes. Our results demonstrate that salamanders have low levels of solo LTRs, suggesting that ectopic-recombination-mediated deletion of LTR retrotransposons occurs more slowly than in other vertebrates with smaller genomes.
Fitzpatrick, Benjamin M; Shaffer, H Bradley
Hybridization between differentiated lineages can have many different consequences depending on fitness variation among hybrid offspring. When introduced organisms hybridize with natives, the ensuing evolutionary dynamics may substantially complicate conservation decisions. Understanding the fitness consequences of hybridization is an important first step in predicting its evolutionary outcome and conservation impact. Here, we measured natural selection caused by differential viability of hybrid larvae in wild populations where native California Tiger Salamanders (Ambystoma californiense) and introduced Barred Tiger Salamanders (Ambystoma tigrinum mavortium) have been hybridizing for 50-60 years. We found strong evidence of hybrid vigor; mixed-ancestry genotypes had higher survival rates than genotypes containing mostly native or mostly introduced alleles. Hybrid vigor may be caused by heterozygote advantage (overdominance) or recombinant hybrid vigor (due to epistasis or complementation). These genetic mechanisms are not mutually exclusive, and we find statistical support for both overdominant and recombinant contributions to hybrid vigor in larval tiger salamanders. Because recombinant homozygous genotypes can breed true, a single highly fit genotype with a mosaic of native and introduced alleles may eventually replace the historically pure California Tiger Salamander (listed as Threatened under the U.S. Endangered Species Act). The management implications of this outcome are complex: Genetically pure populations may not persist into the future, but average fitness and population viability of admixed California Tiger Salamanders may be enhanced. The ecological consequences for other native species are unknown.
Kinkead, K.E.; Otis, D.L.
It has long been accepted that amphibians can skip breeding in any given year, and environmental conditions act as a cue for breeding. In this paper, we quantify temporary emigration or nonbreeding probability for mole and spotted salamanders (Ambystoma talpoideum and A. maculatum). We estimated that 70% of mole salamanders may skip breeding during an average rainfall year and 90% may skip during a drought year. Spotted salamanders may be more likely to breed, with only 17% avoiding the breeding pond during an average rainfall year. We illustrate how superpopulations can be estimated using temporary emigration probability estimates. The superpopulation is the total number of salamanders associated with a given breeding pond. Although most salamanders stay within a certain distance of a breeding pond for the majority of their life spans, it is difficult to determine true overall population sizes for a given site if animals are only captured during a brief time frame each year with some animals unavailable for capture at any time during a given year. ?? 2007 by The Herpetologists' League, Inc.
Kleinteich, Thomas; Herzen, Julia; Beckmann, Felix; Matsui, Masafumi; Haas, Alexander
Larval salamanders (Lissamphibia: Caudata) are known to be effective suction feeders in their aquatic environments, although they will eventually transform into terrestrial tongue feeding adults during metamorphosis. Early tetrapods may have had a similar biphasic life cycle and this makes larval salamanders a particularly interesting model to study the anatomy, function, development, and evolution of the feeding apparatus in terrestrial vertebrates. Here, we provide a description of the muscles that are involved in the feeding strike in salamander larvae of the Hynobiidae and compare them to larvae of the paedomorphic Cryptobranchidae. We provide a functional and evolutionary interpretation for the observed muscle characters. The cranial muscles in larvae from species of the Hynobiidae and Cryptobranchidae are generally very similar. Most notable are the differences in the presence of the m. hyomandibularis, a muscle that connects the hyobranchial apparatus with the lower jaw. We found this muscle only in Onychodactylus japonicus (Hynobiidae) but not in other hynobiid or cryptobranchid salamanders. Interestingly, the m. hyomandibularis in O. japonicus originates from the ceratobranchial I and not the ceratohyal, and thus exhibits what was previously assumed to be the derived condition. Finally, we applied a biomechanical model to simulate suction feeding in larval salamanders. We provide evidence that a flattened shape of the hyobranchial apparatus in its resting position is beneficial for a fast and successful suction feeding strike.
Tyler, T.; Liss, W.J.; Ganio, L.; Larson, Gary L.; Hoffman, Robert L.; Deimling, E.; Lomnicky, G.A.
The larval stage of the long-toed salamander (Ambystoma macrodactylum) is the top vertebrate predator in high-elevation fishless lakes in the North Cascades National Park Service Complex, Washington (U.S.A.). Although most of these high-elevation lakes were naturally fishless, trout have been stocked in many of them. We sought to determine the effects of physicochemical factors and introduced trout on abundance and behavior of A. macrodactylum larvae. Larval salamander densities were estimated by snorkeling. Snorkelers carefully searched through substrate materials within 2 m of the shoreline and recorded the number of larvae observed and if larvae were hidden in benthic substrates. Physicochemical factors were measured in each lake on the same day that snorkel surveys were conducted. In fishless lakes, larval salamander densities were positively related to total Kjeldahl-N concentration and negatively related to lake elevation. Crustacean zooplankton, especially cladocerans, were important food resources for larval A. macrodactylum. Crustacean zooplankton and cladoceran densities were positively related to total Kjeldahl-N, suggesting that increased food resources contributed to increased densities of larval A. macrodactylum. Differences in larval salamander densities between fish and fishless lakes were related to total Kjeldahl-N concentrations and the reproductive status of trout. Mean larval salamander densities for fishless lakes with total Kjeldahl-N amphibians requires an understanding of natural abiotic and biotic factors and processes influencing amphibian distribution and abundance.
Welsh, H.H.; Droege, S.
Terrestrial salamanders of the family P!ethodontidae have unique attributes that make them excellent indicators of biodiversity and ecosystem integrity in forested habitats. Their longevity, small territory size, site fidelity, sensitivity to natural and anthropogenic perturbations, tendency to occur in high densities, and low sampling costs mean that counts of plethodontid salamanders provide numerous advantages over counts of other North American forest organisms for indicating environmental change. Furthermore, they are tightly linked physiologically to microclimatic and successional processes that influence the distribution and abundance of numerous other hydrophilic but difficult-to-study forest-dwelling plants and animals. Ecosystem processes such as moisture cycling, food-web dynamics, and succession, with their related structural and microclimatic variability, all affect forest biodiversity and have been shown to affect salamander populations as well. We determined the variability associated with sampling for plethodontid salamanders by estimating the coefficient of variation (CV) from available time-series data. The median coefficient of variation indicated that variation in counts of individuals among studies was much lower in plethodonticis (27%) than in lepidoptera (93%), passerine birds (57%), small mammals (69%), or other amphibians (37-46%), which means plethodontid salamanders provide an important statistical advantage over other species for monitoring long-term forest health.
Chong, Rebecca A; Mueller, Rachel Lockridge
Mitochondria are the site for the citric acid cycle and oxidative phosphorylation (OXPHOS), the final steps of ATP synthesis via cellular respiration. Each mitochondrion contains its own genome; in vertebrates, this is a small, circular DNA molecule that encodes 13 subunits of the multiprotein OXPHOS electron transport complexes. Vertebrate lineages vary dramatically in metabolic rates; thus, functional constraints on mitochondrial-encoded proteins likely differ, potentially impacting mitochondrial genome evolution. Here, we examine mitochondrial genome evolution in salamanders, which have the lowest metabolic requirements among tetrapods. We show that salamanders experience weaker purifying selection on protein-coding sequences than do frogs, a comparable amphibian clade with higher metabolic rates. In contrast, we find no evidence for weaker selection against mitochondrial genome expansion in salamanders. Together, these results suggest that different aspects of mitochondrial genome evolution (i.e., nucleotide substitution, accumulation of noncoding sequences) are differently affected by metabolic variation across tetrapod lineages.
Cudmore, Wynn W.; Bury, R. Bruce
We investigated the potential for resource partitioning between the Coastal giant salamander (Dicamptodon tenebrosus) and the Cascade torrent salamander (Rhyacotriton cascadae) by examining their diet and microhabitats in forest streams. Larval D. tenebrosus and R. cascadae fed primarily upon aquatic insect larvae. We found similar foods in larval and adult R. cascadae and combined these results. Dicamptodon larvae consumed ephemeropteran, plecopteran, and trichopteran larvae in about equal amounts whereas R. cascadae ate more trichopteran and less ephemeropteran larvae than D. tenebrosus. Diet of all R. cascadae overlapped more with smaller than larger sized D. tenebrosus larvae. Comparisons of diets with available foods indicated R. cascadae is more selective or more gape-limited in its feeding habits than D. tenebrosus larvae. The two salamanders differed in use of microhabitats in creeks, which may contribute to their diet differences.
Martel, A; Blooi, M; Adriaensen, C; Van Rooij, P; Beukema, W; Fisher, M C; Farrer, R A; Schmidt, B R; Tobler, U; Goka, K; Lips, K R; Muletz, C; Zamudio, K R; Bosch, J; Lötters, S; Wombwell, E; Garner, T W J; Cunningham, A A; Spitzen-van der Sluijs, A; Salvidio, S; Ducatelle, R; Nishikawa, K; Nguyen, T T; Kolby, J E; Van Bocxlaer, I; Bossuyt, F; Pasmans, F
Emerging infectious diseases are reducing biodiversity on a global scale. Recently, the emergence of the chytrid fungus Batrachochytrium salamandrivorans resulted in rapid declines in populations of European fire salamanders. Here, we screened more than 5000 amphibians from across four continents and combined experimental assessment of pathogenicity with phylogenetic methods to estimate the threat that this infection poses to amphibian diversity. Results show that B. salamandrivorans is restricted to, but highly pathogenic for, salamanders and newts (Urodela). The pathogen likely originated and remained in coexistence with a clade of salamander hosts for millions of years in Asia. As a result of globalization and lack of biosecurity, it has recently been introduced into naïve European amphibian populations, where it is currently causing biodiversity loss.
Acevedo, Aldemar A; Wake, David B; Márquez, Roberto; Silva, Karen; Franco, Rosmery; Amézquita, Adolfo
The salamander fauna of Colombia is very poorly known, probably because most research efforts have been devoted to anurans during the last two decades. Here, we describe two new species of the genus Bolitoglossa (Eladinea) from the eastern flank of the Eastern Colombian Andes (Cordillera Oriental), near the border with Venezuela. Bolitoglossa tamaense sp. nov. is distributed between 2000 to 2700 m.a.s.l. and Bolitoglossa leandrae sp. nov. is distributed in the low-lands at about 600 m. The new species are diagnosed by a combination of molecular (16S rRNA sequences), coloration, body size, and morphometric (number of maxillary and vomerine teeth and differences in foot webbing) characters. Both species face threats such as chytridiomycosis infections and habitat fragmentation that have already affected other sala-manders in the country. Thus, intensive field and museum work is needed to better document and perhaps protect the local salamander diversity.
Kumar, Anoop; Gates, Phillip B; Czarkwiani, Anna; Brockes, Jeremy P
Limb development in salamanders differs from other tetrapods in that the first digits to form are the two most anterior (preaxial dominance). This has been proposed as a salamander novelty and its mechanistic basis is unknown. Salamanders are the only adult tetrapods able to regenerate the limb, and the contribution of preaxial dominance to limb regeneration is unclear. Here we show that during early outgrowth of the limb bud, a small cohort of cells express the orphan gene Prod1 together with Bmp2, a critical player in digit condensation in amniotes. Disruption of Prod1 with a gene-editing nuclease abrogates these cells, and blocks formation of the radius and ulna, and outgrowth of the anterior digits. Preaxial dominance is a notable feature of limb regeneration in the larval newt, but this changes abruptly after metamorphosis so that the formation of anterior and posterior digits occurs together within the autopodium resembling an amniote-like pattern.
Morrison, Jamie I; Lööf, Sara; He, Pingping; Simon, András
In contrast to mammals, salamanders can regenerate complex structures after injury, including entire limbs. A central question is whether the generation of progenitor cells during limb regeneration and mammalian tissue repair occur via separate or overlapping mechanisms. Limb regeneration depends on the formation of a blastema, from which the new appendage develops. Dedifferentiation of stump tissues, such as skeletal muscle, precedes blastema formation, but it was not known whether dedifferentiation involves stem cell activation. We describe a multipotent Pax7+ satellite cell population located within the skeletal muscle of the salamander limb. We demonstrate that skeletal muscle dedifferentiation involves satellite cell activation and that these cells can contribute to new limb tissues. Activation of salamander satellite cells occurs in an analogous manner to how the mammalian myofiber mobilizes stem cells during skeletal muscle tissue repair. Thus, limb regeneration and mammalian tissue repair share common cellular and molecular programs. Our findings also identify satellite cells as potential targets in promoting mammalian blastema formation.
Sandoval-Guzmán, Tatiana; Wang, Heng; Khattak, Shahryar; Schuez, Maritta; Roensch, Kathleen; Nacu, Eugeniu; Tazaki, Akira; Joven, Alberto; Tanaka, Elly M; Simon, András
Salamanders regenerate appendages via a progenitor pool called the blastema. The cellular mechanisms underlying regeneration of muscle have been much debated but have remained unclear. Here we applied Cre-loxP genetic fate mapping to skeletal muscle during limb regeneration in two salamander species, Notophthalmus viridescens (newt) and Ambystoma mexicanum (axolotl). Remarkably, we found that myofiber dedifferentiation is an integral part of limb regeneration in the newt, but not in axolotl. In the newt, myofiber fragmentation results in proliferating, PAX7(-) mononuclear cells in the blastema that give rise to the skeletal muscle in the new limb. In contrast, myofibers in axolotl do not generate proliferating cells, and do not contribute to newly regenerated muscle; instead, resident PAX7(+) cells provide the regeneration activity. Our results therefore show significant diversity in limb muscle regeneration mechanisms among salamanders and suggest that multiple strategies may be feasible for inducing regeneration in other species, including mammals.
Full Text Available The Chinese giant salamander belongs to an old lineage of salamanders and endangered species. Many studies of breeding and disease regarding this amphibian had been implemented. However, the studies on the ultrastructure of this amphibian are rare. In this work, we provide a histological and ultrastructural investigation on posterior esophagus of Chinese giant salamander. The sections of amphibian esophagus were stained by hematoxylin & eosin (H&E. Moreover, the esophageal epithelium was observed by transmission electron microscopy (TEM. The results showed that esophageal epithelium was a single layer epithelium, which consisted of mucous cells and columnar cells. The esophageal glands were present in submucosa. The columnar cells were ciliated. According to the diverging ultrastructure of mucous vesicles, three types of mucous cells could be identified in the esophageal mucosa: i electron-lucent vesicles mucous cell (ELV-MC; ii electron-dense vesicles mucous cell (EDV-MC; and iii mixed vesicles mucous cell (MV-MC.
Sebastiano, Salvidio; Antonio, Romano; Fabrizio, Oneto; Dario, Ottonello; Roberta, Michelon
Trophic niche may be the most important ecological dimension for some vertebrate groups and in particular for terrestrial amphibians, that are important predators of soil invertebrates. In general, resource partitioning occurs between syntopic species with similar ecological niches, and coexistence patterns seem to be regulated by temporal resource variability. However most of the generalization on foraging strategies of terrestrial salamanders are extrapolated from studies on New World temperate species, thus we investigated the seasonal effect of resource variation in an European forest ecosystem, in which two ecologically similar but phylogenetically distinct salamander species are found. The diet of adult and juvenile cave salamanders (Speleomantes strinati), and of adult spectacled salamander (Salamandrina perspicillata) was obtained by stomach flushing, and results showed large seasonal changes both in prey availability and in salamander realised trophic niche. Values of trophic diversity were similar and niche overlaps were large among all salamander groups in spring, during high prey availability. Conversely in autumn, when a two-fold reduction in prey biomass was observed, there was a clear niche partitioning as the smaller S. perspicillata shifted from a generalist to a specialized trophic strategy. Juvenile Speleomantes strinatii, that largely overlapped in size with S. perspicillata, did not show any change in diet, suggesting that the feeding strategies were species-specific and not size-mediated. The observed patterns of variation in feeding ecology indicate that similar predators may react differently to changing prey availability to enhance niche partitioning. We also observed an increased energy intake during autumn for S perspicillata and S. strinatii juveniles, possibly related to differences in microhabitat use and activity patterns.
Charrier, V; Cabelguen, J-M
Most investigations into the role of the body axis in vertebrate locomotion have focused on the trunk, although in most tetrapods, the tail also plays an active role. In salamanders, the tail contributes to propulsion during swimming and to dynamic balance and maneuverability during terrestrial locomotion. The aim of the present study was to obtain information concerning the neural mechanisms that produce tail muscle contractions during locomotion in the salamander Pleurodeles waltlii. We recorded the ventral root activities in in vitro spinal cord preparations in which locomotor-like activity was induced via bath application of N-methyl-d-aspartate (20μM) and d-serine (10μM). Recordings showed that the tail spinal cord is capable of producing propagated waves of motor activity that alternate between the left and right sides. Lesion experiments further revealed that the tail rhythmogenic network is composed of a double chain of identical hemisegmental oscillators. Finally, using spinal cord preparations bathed in a chamber partitioned into two pools, we revealed efficient short-distance coupling between the trunk and tail networks. Together, our results demonstrate the existence of a pattern generator for rhythmic tail movements in the salamander and show that the global architecture of the tail network is similar to that previously proposed for the mid-trunk locomotor network in the salamander. Our findings further support the view that salamanders can control their trunk and tail independently during stepping movements. The relevance of our results in relation to the generation of tail muscle contractions in freely moving salamanders is discussed.
Laking, Alexandra E.; Ngo, Hai Ngoc; Pasmans, Frank; Martel, An; Nguyen, Tao Thien
The amphibian chytrid fungi, Batrachochytrium dendrobatidis (Bd) and B. salamandrivorans (Bsal), pose a major threat to amphibian biodiversity. Recent evidence suggests Southeast Asia as a potential cradle for both fungi, which likely resulted in widespread host-pathogen co-existence. We sampled 583 salamanders from 8 species across Vietnam in 55 locations for Bsal and Bd, determined scaled mass index as a proxy for fitness and collected environmental data. Bsal was found within 14 of the 55 habitats (2 of which it was detected in 2013), in 5 salamandrid species, with a prevalence of 2.92%. The globalized pandemic lineage of Bd was found within one pond on one species with a prevalence of 0.69%. Combined with a complete lack of correlation between infection and individual body condition and absence of indication of associated disease, this suggests low level pathogen endemism and Bsal and Bd co-existence with Vietnamese salamandrid populations. Bsal was more widespread than Bd, and occurs at temperatures higher than tolerated by the type strain, suggesting a wider thermal niche than currently known. Therefore, this study provides support for the hypothesis that these chytrid fungi may be endemic to Asia and that species within this region may act as a disease reservoir. PMID:28287614
Trumbo, Daryl R; Spear, Stephen F; Baumsteiger, Jason; Storfer, Andrew
A species' genetic structure often varies in response to ecological and landscape processes that differ throughout the species' geographic range, yet landscape genetics studies are rarely spatially replicated. The Cope's giant salamander (Dicamptodon copei) is a neotenic, dispersal-limited amphibian with a restricted geographic range in the Pacific northwestern USA. We investigated which landscape factors affect D. copei gene flow in three regions spanning the species' range, which vary in climate, landcover and degree of anthropogenic disturbance. Least cost paths and Circuitscape resistance analyses revealed that gene flow patterns vary across the species' range, with unique combinations of landscape variables affecting gene flow in different regions. Populations in the northern coastal portions of the range had relatively high gene flow, largely facilitated by stream and river networks. Near the southeastern edge of the species' range, gene flow was more restricted overall, with relatively less facilitation by streams and more limitation by heat load index and fragmented forest cover. These results suggested that the landscape is more difficult for individuals to disperse through at the southeastern edge of the species' range, with terrestrial habitat desiccation factors becoming more limiting to gene flow. We suggest that caution be used when attempting to extrapolate landscape genetic models and conservation measures from one portion of a species' range to another.
Full Text Available Populations of Mexican plethodontid salamanders have been surveyed non-systematically over the last 25 years. In light of many reports of disappearance of amphibians around the world, we checked for persistence of reported species at ten of these sites. All of the commoner species persist (we observed individuals representing a total of 30 species. While observed densities of many species of Mexican plethodontids are lower to much lower than was the case 20 to 25 years ago, evidence for recent extinctions, such as has been reported for amphibian taxa elsewhere, is equivocal or lacking. Habitat modification has contributed to difficulties in finding certain species.Poblaciones de varias especies de salamandras pletodóntidas en México han sido monitoreadas de manera no sistemática durante los últimos 25 años. Diez de éstas poblaciones fueran visitadas recientemente con el propósito de verificar la persistencia de las especies reportadas para dichas localidades. Nuestras observaciones confirman la persistencia local de más de 30 especies cuyo estatus era desconocido, aunque la frecuencia de observación de estas especies es en general menor que en fechas anteriores. Estas observaciones son particularmente relevantes dada la situación actual de preocupación por la disminución mundial de anfibios.
Yin, Xin-Yan; Jia, Li-Chao; Wang, Chen; Xie, Guang-Ming
Amphibious salamanders often swing their waist to coordinate quadruped walking in order to improve their crawling speed. A robot with a swing waist joint, like an amphibious salamander, is used to mimic this locomotion. A control method is designed to allow the robot to maintain the rotational speed of its legs continuous and avoid impact between its legs and the ground. An analytical expression is established between the amplitude of the waist joint and the step length. Further, an optimization amplitude is obtained corresponding to the maximum stride. The simulation results based on automatic dynamic analysis of mechanical systems (ADAMS) and physical experiments verify the rationality and validity of this expression.
Xin-Yan Yin; Li-Chao Jia; Chen Wang; Guang-Ming Xie
Amphibious salamanders often swing their waist to coordinate quadruped walking in order to improve their crawling speed. A robot with a swing waist joint, like an amphibious salamander, is used to mimic this locomotion. A control method is designed to allow the robot to maintain the rotational speed of its legs continuous and avoid impact between its legs and the ground. An analytical expression is established between the amplitude of the waist joint and the step length. Further, an optimization amplitude is obtained corresponding to the maximum stride. The simulation results based on automatic dynamic analysis of mechanical systems (ADAMS) and physical experiments verify the rationality and validity of this expression.
Chang, Jin-Soo; Gu, Man Bock; Kim, Kyoung-Woong
The p53 mutation in salamanders can be used as an indicator of arsenic contamination. The influence of arsenic exposure was studied on mutation of tumor suppressor gene in salamanders collected from several As-contaminated mine areas in Korea. Salamander eggs and larvae were exposed to arsenic in a toxicity test, and teratogenic salamanders found in heavy metal- and As-contaminated water from As-Bi mines were evaluated using PCR-SSCP to determine if they would be useful as an ecological indicator species. Changes in amino acids were shown to have occurred as a result of an arsenic-accumulating event that occurred after the DNA damage. In addition, both of the Hynobius leechii exposed groups were primarily affected by forms of skin damage, changes in the lateral tail/dorsal flexure and/or abnormality teratogenesis. Single-base sense mutation in codons 346 (AAG: Lys to ATG: Met), 224 (TTT: Phe to TTA: Leu), 211 (ATG: Met to AAG: Lys), 244 (TTT: Phe to TTTG: insertion), 245 (Glu GAG to Gln CAG) and 249 (TGT Cys to TGA stop) of the p53 gene were simultaneously found in mutated salamanders. Based on the results of our data illustrating the effect of arsenic exposure on the p53 mutation of salamanders in arsenic-contaminated mine areas, these mutated salamanders can be used as potential ecological indicators in the arsenic-contaminated ecosystems.
Skutschas, Pavel; Stein, Koen
Kokartus honorarius from the Middle Jurassic (Bathonian) of Kyrgyzstan is one of the oldest salamanders in the fossil record, characterized by a mixture of plesiomorphic morphological features and characters shared with crown-group salamanders. Here we present a detailed histological analysis of its long bones. The analysis of a growth series demonstrates a significant histological maturation during ontogeny, expressed by the progressive appearance of longitudinally oriented primary vascular canals, primary osteons, growth marks, remodelling features in primary bone tissues, as well as progressive resorption of the calcified cartilage, formation of endochondral bone and development of cartilaginous to bony trabeculae in the epiphyses. Apart from the presence of secondary osteons, the long bone histology of Kokartus is very similar to that of miniaturized temnospondyls, other Jurassic stem salamanders, miniaturized seymouriamorphs and modern crown-group salamanders. We propose that the presence of secondary osteons in Kokartus honorarius is a plesiomorphic feature, and the loss of secondary osteons in the long bones of crown-group salamanders as well as in those of miniaturized temnospondyls is the result of miniaturization processes. Hitherto, all stem salamander long bong histology (Kokartus, Marmorerpeton and 'salamander A') has been generally described as having paedomorphic features (i.e. the presence of Katschenko's Line and a layer of calcified cartilage), these taxa were thus most likely neotenic forms. The absence of clear lines of arrested growth and annuli in long bones of Kokartus honorarius suggests that the animals lived in an environment with stable local conditions.
little is known about the ecology of the chinese giant salamander (andrias davidianus),a critically endangered species.such information is needed to make informed decisions concerning the conservation and management of this species.four a.davidianus raised in a pool were released into their native habitat on 04 may 2005 and were subsequently radio-tracked for approximately 155-168 days.following their release,the giant salamanders traveled upstream in search of suitable micro-habitats,and settled after 10 days.later,a devastating summer flash flood destroyed the salamanders' dens,triggering another bout of habitat searching by the animals.eventually,the salamanders settled in different sections of the stream where they remained until the end of the study.on average,each habitat searching endeavor took 7.5 days,during which a giant salamander explored a 310 m stretch of stream with a surface area of about 1157 m2 and occupied 3.5 temporary dwellings.each giant salamander spent an average of 144.5 days in semi-permanent micro-habitats,and occupied territories that had a mean size of 34.75 m2.our results indicate that the chinese giant salamander responds to habitat disturbance by seeking new habitats upstream,both water temperature and water level affect the salamander's habitat searching activity,and the size of the salamander's semi-permanent territory is influenced by the size of the pool containing the animal's den.
Walker, Donald M; Leys, Jacob E; Dunham, Kelly E; Oliver, Joshua C; Schiller, Emily E; Stephenson, Kelsey S; Kimrey, John T; Wooten, Jessica; Rogers, Mark W
Environmental DNA (eDNA) can be used as an assessment tool to detect populations of threatened species and provide fine-scale data required to make management decisions. The objectives of this project were to use quantitative PCR (qPCR) to: (i) detect spiked salamander DNA in soil, (ii) quantify eDNA degradation over time, (iii) determine detectability of salamander eDNA in a terrestrial environment using soil, faeces, and skin swabs, (iv) detect salamander eDNA in a mesocosm experiment. Salamander eDNA was positively detected in 100% of skin swabs and 66% of faecal samples and concentrations did not differ between the two sources. However, eDNA was not detected in soil samples collected from directly underneath wild-caught living salamanders. Salamander genomic DNA (gDNA) was detected in all qPCR reactions when spiked into soil at 10.0, 5.0, and 1.0 ng/g soil and spike concentration had a significant effect on detected concentrations. Only 33% of samples showed recoverable eDNA when spiked with 0.25 ng/g soil, which was the low end of eDNA detection. To determine the rate of eDNA degradation, gDNA (1 ng/g soil) was spiked into soil and quantified over seven days. Salamander eDNA concentrations decreased across days, but eDNA was still amplifiable at day 7. Salamander eDNA was detected in two of 182 mesocosm soil samples over 12 weeks (n = 52 control samples; n = 65 presence samples; n = 65 eviction samples). The discrepancy in detection success between experiments indicates the potential challenges for this method to be used as a monitoring technique for small-bodied wild terrestrial salamander populations. © 2017 John Wiley & Sons Ltd.
Effects of Buffering Key Habitat for Terrestrial Salamanders: Implications for the Management of the Federally Threatened Red Hills Salamander (Phaeognathus hubrichti and Other Imperiled Plethodontids
Joseph J. Apodaca
Full Text Available Forestry practices are placing ever increasing emphasis on sustainability and the maintenance of ecological processes, biodiversity, and endangered species or populations. Balancing timber harvest and the management of imperiled species presents a particularly difficult challenge during this shift, as we often know very little about these species’ natural history and how and why silviculture practices affect their populations. Accordingly, investigation of and improvement on current management practices for threatened species is imperative. We investigated the effectiveness of habitat buffers as a management technique for the imperiled Red Hills salamander (Phaeognathus hubrichti by combining genetic, transect, and body-condition data. We found that populations where habitat buffers have been employed have higher genetic diversity and higher population densities, and individuals have better overall body condition. These results indicate that buffering the habitat of imperiled species can be an effective management tool for terrestrial salamanders. Additionally, they provide further evidence that leaving the habitat of imperiled salamanders unbuffered can have both immediate and long-term negative impacts on populations.
Stark, Tariq; Laurijssens, Carlijn; Weterings, Martijn; Martel, An; Köhler, Gunther; Pasmans, Frank
Amphibians are the most threatened terrestrial vertebrates on the planet and are iconic in the global biodiversity crisis. Their global decline caused by the fungal agent Batrachochytrium dendrobatidis (Bd) is well known. Declines of Mesoamerican salamanders of the family Plethodontidae, mainly affe
Andrés R. Acosta-Galvis
Full Text Available Eight species of salamanders are recognized to Cordillera Oriental of Colombia. Here we describe a new species of the genus Bolitoglossa, named Bolitoglossa guaneae sp. nov. The highest number of species of this genus is found in the cloud forests located in the western flank of the Cordillera Oriental.
Sepulveda, Adam J; Lowe, Winsor H
Theory suggests that source-sink dynamics can allow coexistence of intraguild predators and prey, but empirical evidence for this coexistence mechanism is limited. We used capture-mark-recapture, genetic methods, and stable isotopes to test whether source-sink dynamics promote coexistence between stream fishes, the intraguild predator, and stream salamanders (Dicamptodon aterrimus), the intraguild prey. Salamander populations from upstream reaches without fish were predicted to maintain or supplement sink populations in downstream reaches with fish. We found instead that downstream reaches with fish were not sinks even though fish consumed salamander larvae-apparent survival, recruitment, and population growth rate did not differ between upstream and downstream reaches. There was also no difference between upstream and downstream reaches in net emigration. We did find that D. aterrimus moved frequently along streams, but believe that this is a response to seasonal habitat changes rather than intraguild predation. Our study provides empirical evidence that local-scale mechanisms are more important than dispersal dynamics to coexistence of streams salamanders and fish. More broadly, it shows the value of empirical data on dispersal and gene flow for distinguishing between local and spatial mechanisms of coexistence.
Chatfield, Matthew W H; Moler, Paul; Richards-Zawacki, Corinne L
Little is known about the impact that the pathogenic amphibian chytrid fungus, Batrachochytrium dendrobatidis (Bd), has on fully aquatic salamander species of the eastern United States. As a first step in determining the impacts of Bd on these species, we aimed to determine the prevalence of Bd in wild populations of fully aquatic salamanders in the genera Amphiuma, Necturus, Pseudobranchus, and Siren. We sampled a total of 98 salamanders, representing nine species from sites in Florida, Mississippi, and Louisiana. Overall, infection prevalence was found to be 0.34, with significant differences among genera but no clear geographic pattern. We also found evidence for seasonal variation, but additional sampling throughout the year is needed to clarify this pattern. The high rate of infection discovered in this study is consistent with studies of other amphibians from the southeastern United States. Coupled with previously published data on life histories and population densities, the results presented here suggest that fully aquatic salamanders may be serving as important vectors of Bd and the interaction between these species and Bd warrants additional research.
US Fish and Wildlife Service, Department of the Interior — The objective of the survey was to document Cheat Mountain salamander use on either side of Powderline ski trail or Three-Mile ski trail in an effort to continue to...
NIR spectra were collected at three surface locations for Chinese giant salamanders to ascertain whether spectral signatures could be separated by anatomical, presumably physiologically-based, locations. The first location was the smooth area immediately above the cloaca on the animal’s abdomen, whi...
Greenwald, Katherine R; Lisle Gibbs, H
Unisexual (all female) salamanders in the genus Ambystoma are animals of variable ploidy (2N-5N) that reproduce via a unique system of 'leaky' gynogenesis. As a result, these salamanders have a diverse array of nuclear genome combinations from up to five sexual species: the blue-spotted (A. laterale), Jefferson (A. jeffersonianum), smallmouth (A. texanum), tiger (A. tigrinum) and streamside (A. barbouri) salamanders. Identifying the genome complement, or biotype, is a critical first step in addressing a broad range of ecological and evolutionary questions about these salamanders. Previous work relied upon genome-related differences in allele size distributions for specific microsatellite loci, but overlap in these distributions among different genomes makes definitive identification and ploidy determination in unisexuals difficult or impossible. Here, we develop the first single nucleotide polymorphism assay for the identification of unisexual biotypes, based on species-specific nucleotide polymorphisms in noncoding DNA loci. Tests with simulated and natural unisexual DNA samples show that this method can accurately identify genome complement and estimate ploidy, making this a valuable tool for assessing the genome composition of unisexual samples.
Chudyk, Sarah; McMillan, Amy; Lange, Catherine
This article contains an original 5E lesson plan developed from conservation genetics research on the giant North American hellbender salamander, Cryptobranchus alleganiensis alleganiensis. The lesson plan provides background information on the hellbender, reviews basic genetics, and exposes students to the scientific process that is used during…
Thien, Tao Nguyen; Martel, An; Brutyn, Melanie; Bogaerts, Sergé; Sparreboom, Max; Haesebrouck, Freddy; Fisher, Matthew C; Beukema, Wouter; Van, Tang Duong; Chiers, Koen; Pasmans, Frank
Until now, Asian amphibians appear to have largely escaped declines driven by chytridiomycosis. Vietnamese salamanders that belong to the genus Tylototriton are rare and have a patchy distribution in mountainous areas, falling within the proposed environmental envelope of chytrid infections, surrounded by Batrachochytrium dendrobatidis infected regions. If these salamanders are susceptible to chytridiomycosis, then their populations could be highly vulnerable after the introduction of B. dendrobatidis. Examination for the presence of the chytrid fungus in skin swabs from 19 Tylototriton asperrimus and 104 Tylototriton vietnamensis by using quantitative polymerase chain reaction was performed. Susceptibility of T. asperrimus to experimental infection by using the global panzootic lineage (BdGPL) strain of B. dendrobatidis was examined. The fungus was absent in all samples from all wild salamanders examined. Inoculation with the BdGPL strain resulted in mortality of all five inoculated salamanders within 3 weeks after inoculation with infected animals that manifested severe orthokeratotic hyperkeratosis, epidermal hyperplasia, and spongiosis. Although infection by B. dendrobatidis currently appears absent in Vietnamese Tylototriton populations, the rarity of these animals, their pronounced susceptibility to chytridiomycosis, an apparently suitable environmental context and increasing likelihood of the pathogen being introduced, together suggest the need of urgent measures to avoid future scenarios of extinction as witnessed in Central America and Australia.
Abdullayev, Ilgar; Kirkham, Matthew; Björklund, Åsa K; Simon, András; Sandberg, Rickard
Salamanders have a remarkable capacity to regenerate complex tissues, such as limbs and brain, and are therefore an important comparative model system for regenerative medicine. Despite these unique properties among adult vertebrates, the genomic information for amphibians in general, and salamanders in particular, is scarce. Here, we used massive parallel sequencing to reconstruct a de novo reference transcriptome of the red spotted newt (Notophthalmus viridescens) containing 118,893 transcripts with a N50 length of 2016 nts. Comparisons to other vertebrates revealed a newt transcriptome that is comparable in size and characteristics to well-annotated vertebrate transcriptomes. Identification of putative open reading frames (ORFs) enabled us to infer a comprehensive proteome, including the annotation of 19,903 newt proteins. We used the identified domain architectures (DAs) to assign ORFs phylogenetic positions, which also revealed putative salamander specific proteins. The reference transcriptome and inferred proteome of the red spotted newt will facilitate the use of systematic genomic technologies for regeneration studies in salamanders and enable evolutionary analyses of vertebrate regeneration at the molecular level.
Weisrock David W
Full Text Available Abstract Salamanders of the genus Ambystoma are a unique model organism system because they enable natural history and biomedical research in the laboratory or field. We developed Sal-Site to integrate new and existing ambystomatid salamander research resources in support of this model system. Sal-Site hosts six important resources: 1 Salamander Genome Project: an information-based web-site describing progress in genome resource development, 2 Ambystoma EST Database: a database of manually edited and analyzed contigs assembled from ESTs that were collected from A. tigrinum tigrinum and A. mexicanum, 3 Ambystoma Gene Collection: a database containing full-length protein-coding sequences, 4 Ambystoma Map and Marker Collection: an image and database resource that shows the location of mapped markers on linkage groups, provides information about markers, and provides integrating links to Ambystoma EST Database and Ambystoma Gene Collection databases, 5 Ambystoma Genetic Stock Center: a website and collection of databases that describe an NSF funded salamander rearing facility that generates and distributes biological materials to researchers and educators throughout the world, and 6 Ambystoma Research Coordination Network: a web-site detailing current research projects and activities involving an international group of researchers. Sal-Site is accessible at http://www.ambystoma.org.
Bendik, Nathan F.; Meik, Jesse M; Gluesenkamp, Andrew G.; Roelke, Corey E; Chippindale, Paul T
Background Subterranean faunal radiations can result in complex patterns of morphological divergence involving both convergent or parallel phenotypic evolution and cryptic species diversity. Salamanders of the genus Eurycea in central Texas provide a particularly challenging example with respect to phylogeny reconstruction, biogeography and taxonomy. These predominantly aquatic species inhabit karst limestone aquifers and spring outflows, and exhibit a wide range of morphological and genetic ...
Hart Welsh; Garth Hodgson
Woodland (Plethodontid) salamanders occur in huge numbers in healthy forests in North America where the abundances of many species vary along successional gradients. Their high numbers and trophic role as predators on shredder and decomposer arthropods influence nutrient and carbon pathways at the leaf litter/soil interface. Their extreme niche conservatism and low...
Full Text Available The Chinese giant salamander (Andrias davidianus, renowned as a living fossil, is the largest and longest-lived amphibian species in the world. Its skin is rich in collagens, and has developed mucous gland which could secrete a large amount of mucus under the scraping and electric stimulation. The molting is the degraded skin stratum corneum. To establish the functional skin proteome of Chinese giant salamander, two-dimensional gel electrophoresis (2DE and mass spectrometry (MS were applied to detect the composition and relative abundance of the proteins in the skin, mucus and molting. The determination of the general proteome in the skin can potentially serve as a foundation for future studies characterizing the skin proteomes from diseased salamander to provide molecular and mechanistic insights into various disease states and potential therapeutic interventions. Data presented here are also related to the research article “Proteomic analysis of the skin of Chinese giant salamander (Andrias davidianus” in the Journal of Proteomics .
Full Text Available Fossils are almost always represented by hard tissues but we present here the exceptional case of a three-dimensionally preserved specimen that was ‘mummified’ (likely between 40 and 34 million years ago in a terrestrial karstic environment. This fossil is the incomplete body of a salamander, Phosphotriton sigei, whose skeleton and external morphology are well preserved, as revealed by phase-contrast synchrotron X-ray microtomography. In addition, internal structures composed of soft tissues preserved in three dimensions are now identified: a lung, the spinal cord, a lumbosacral plexus, the digestive tract, muscles and urogenital organs that may be cloacal glands. These are among the oldest known cases of three-dimensional preservation of these organs in vertebrates and shed light on the ecology of this salamander. Indeed, the digestive tract contains remains of a frog, which represents the only known case of an extinct salamander that fed on a frog, an extremely rare type of predation in extant salamanders. These new data improve our scarce knowledge on soft tissue anatomy of early urodeles and should prove useful for future biologists and palaeontologists working on urodele evolutionary biology. We also suggest that the presence of bat guano and carcasses represented a close source of phosphorus, favouring preservation of soft tissues. Bone microanatomy indicates that P. sigei was likely amphibious or terrestrial, and was probably not neotenic.
Jéssica Barata da Silva
Full Text Available Plethodontid salamanders of genus Bolitoglossa constitute the largest and most diverse group of salamanders, including around 20% of living caudate species. Recent studies have indicated the occurrence of five recognized species in the Brazilian Amazon Rainforest. We present here the first cytogenetic data of a Brazilian salamander, which may prove to be a useful by contribution to the cytotaxonomy of the genus. Specimens were collected near the "type" locality (Utinga, Belém, PA, Brazil. Chromosomal preparations from duodenal epithelial cells and testes were subjected to Giemsa staining, C-banding and DAPI/CMA3 fluorochrome staining. All specimens showed a karyotype with 13 bi-armed chromosome pairs (2n = 26. Nucleolar Organizer Regions, evidenced by CMA3, were located distally on the long arm of pair 7 (7q. DAPI+ heterochromatin was predominantly centromeric, with some small pericentromeric bands. Although the C-banding patterns of other Bolitoglossa species are so far unknown, cytogenetic studies conducted in other Plethodontid salamanders have demonstrated that pericentromeric heterochromatin is a useful cytological marker for identifying interspecific homeologies. Species diversification is usually accompanied by chromosomal changes. Therefore, the cytogenetic characterization of Bolitoglossa populations from the middle and western Brazilian Amazon Basin could identify differences which may lead to the identification of new species.
Mueller, Rachel Lockridge; Boore, Jeffrey L.
Extensive gene rearrangement is reported in the mitochondrial genomes of lungless salamanders (Plethodontidae). In each genome with a novel gene order, there is evidence that the rearrangement was mediated by duplication of part of the mitochondrial genome, including the presence of both pseudogenes and additional, presumably functional, copies of duplicated genes. All rearrangement-mediating duplications include either the origin of light strand replication and the nearby tRNA genes or the regions flanking the origin of heavy strand replication. The latter regions comprise nad6, trnE, cob, trnT, an intergenic spacer between trnT and trnP and, in some genomes, trnP, the control region, trnF, rrnS, trnV, rrnL, trnL1, and nad1. In some cases, two copies of duplicated genes, presumptive regulatory regions, and/or sequences with no assignable function have been retained in the genome following the initial duplication; in other genomes, only one of the duplicated copies has been retained. Both tandem and non-tandem duplications are present in these genomes, suggesting different duplication mechanisms. In some of these mtDNAs, up to 25 percent of the total length is composed of tandem duplications of non-coding sequence that includes putative regulatory regions and/or pseudogenes of tRNAs and protein-coding genes along with otherwise unassignable sequences. These data indicate that imprecise initiation and termination of replication, slipped-strand mispairing, and intra-molecular recombination may all have played a role in generating repeats during the evolutionary history of plethodontid mitochondrial genomes.
Background The Mexican axolotl (Ambystoma mexicanum) is considered a hopeful monster because it exhibits an adaptive and derived mode of development - paedomorphosis - that has evolved rapidly and independently among tiger salamanders. Unlike related tiger salamanders that undergo metamorphosis, axolotls retain larval morphological traits into adulthood and thus present an adult body plan that differs dramatically from the ancestral (metamorphic) form. The basis of paedomorphic development was investigated by comparing temporal patterns of gene transcription between axolotl and tiger salamander larvae (Ambystoma tigrinum tigrinum) that typically undergo a metamorphosis. Results Transcript abundances from whole brain and pituitary were estimated via microarray analysis on four different days post hatching (42, 56, 70, 84 dph) and regression modeling was used to independently identify genes that were differentially expressed as a function of time in both species. Collectively, more differentially expressed genes (DEGs) were identified as unique to the axolotl (n = 76) and tiger salamander (n = 292) than were identified as shared (n = 108). All but two of the shared DEGs exhibited the same temporal pattern of expression and the unique genes tended to show greater changes later in the larval period when tiger salamander larvae were undergoing anatomical metamorphosis. A second, complementary analysis that directly compared the expression of 1320 genes between the species identified 409 genes that differed as a function of species or the interaction between time and species. Of these 409 DEGs, 84% exhibited higher abundances in tiger salamander larvae at all sampling times. Conclusions Many of the unique tiger salamander transcriptional responses are probably associated with metamorphic biological processes. However, the axolotl also showed unique patterns of transcription early in development. In particular, the axolotl showed a genome-wide reduction in mRNA abundance
Full Text Available Abstract Background The Mexican axolotl (Ambystoma mexicanum is considered a hopeful monster because it exhibits an adaptive and derived mode of development - paedomorphosis - that has evolved rapidly and independently among tiger salamanders. Unlike related tiger salamanders that undergo metamorphosis, axolotls retain larval morphological traits into adulthood and thus present an adult body plan that differs dramatically from the ancestral (metamorphic form. The basis of paedomorphic development was investigated by comparing temporal patterns of gene transcription between axolotl and tiger salamander larvae (Ambystoma tigrinum tigrinum that typically undergo a metamorphosis. Results Transcript abundances from whole brain and pituitary were estimated via microarray analysis on four different days post hatching (42, 56, 70, 84 dph and regression modeling was used to independently identify genes that were differentially expressed as a function of time in both species. Collectively, more differentially expressed genes (DEGs were identified as unique to the axolotl (n = 76 and tiger salamander (n = 292 than were identified as shared (n = 108. All but two of the shared DEGs exhibited the same temporal pattern of expression and the unique genes tended to show greater changes later in the larval period when tiger salamander larvae were undergoing anatomical metamorphosis. A second, complementary analysis that directly compared the expression of 1320 genes between the species identified 409 genes that differed as a function of species or the interaction between time and species. Of these 409 DEGs, 84% exhibited higher abundances in tiger salamander larvae at all sampling times. Conclusions Many of the unique tiger salamander transcriptional responses are probably associated with metamorphic biological processes. However, the axolotl also showed unique patterns of transcription early in development. In particular, the axolotl showed a genome
The pupils in final year of high school (15-18 years old) study the notion "species" and the creation of new species in various ways. Having studied genetic admixtures, this activity allows the pupils to build a scenario explaining the creation of a new species of Salamander in southern California from an ancestral population existing in northern Oregon. They can observe, on Google Earth, various populations of Salamander of the genus Ensatina. Salamanders of the genus Ensatina live in California around the Joaquin and Sacramento dry valleys. In this software, the pupils get information about the salamanders' environment and photographs of individuals and environments. During a migratory movement toward new territories to be colonized, these salamanders meet an inhospitable environment that they can not occupy. This population then splits up into two migratory branches, east and west, each overcoming the obstacles in different ways. The two groups gradually colonized southern territories but they avoided the too dry and hot San Joaquin plains. The two main branches of the original population gradually move away from each other, and genetic exchanges between them decrease over time. Eventually, we can find various populations of Salamander on both sides of the valleys, since the salamanders occupied new territories and diversified along the way. Among mutations that randomly occur, only those mutations that are best adapted in the origin were conserved in the genetic heritage of every population. When the individuals stemming from different western populations met, they were interfertile and give fertile hybrids, which was verified in the laboratory. Likewise, when individuals of the different eastern subspecies met accidentally, fertile hybrids also could arise from these crossings. The pupils can observe what happens in the overlap of various populations : interfertility or not. They also have geological, geographical and climatic information about the San Joaquin
Andrew J Kroll
Full Text Available Plethodontid salamanders are diverse and widely distributed taxa and play critical roles in ecosystem processes. Due to salamander use of structurally complex habitats, and because only a portion of a population is available for sampling, evaluation of sampling designs and estimators is critical to provide strong inference about Plethodontid ecology and responses to conservation and management activities. We conducted a simulation study to evaluate the effectiveness of multi-scale and hierarchical single-scale occupancy models in the context of a Before-After Control-Impact (BACI experimental design with multiple levels of sampling. Also, we fit the hierarchical single-scale model to empirical data collected for Oregon slender and Ensatina salamanders across two years on 66 forest stands in the Cascade Range, Oregon, USA. All models were fit within a Bayesian framework. Estimator precision in both models improved with increasing numbers of primary and secondary sampling units, underscoring the potential gains accrued when adding secondary sampling units. Both models showed evidence of estimator bias at low detection probabilities and low sample sizes; this problem was particularly acute for the multi-scale model. Our results suggested that sufficient sample sizes at both the primary and secondary sampling levels could ameliorate this issue. Empirical data indicated Oregon slender salamander occupancy was associated strongly with the amount of coarse woody debris (posterior mean = 0.74; SD = 0.24; Ensatina occupancy was not associated with amount of coarse woody debris (posterior mean = -0.01; SD = 0.29. Our simulation results indicate that either model is suitable for use in an experimental study of Plethodontid salamanders provided that sample sizes are sufficiently large. However, hierarchical single-scale and multi-scale models describe different processes and estimate different parameters. As a result, we recommend careful consideration of
Bank, M. S.; Crocker, J.; Wachtl, J.; Kleeman, P.; Fellers, G.; Currens, C.; Hothem, R.; Madej, M. A.
Mercury (Hg) contamination of stream salamanders in the Pacific Northwest region of the United States has received little attention. Here we report total Hg (HgT) and methyl mercury (MeHg) concentrations in larval giant salamanders (Dicamptodon spp.) and surface water from forested and chaparral lotic ecosystems distributed along a latitudinal gradient throughout Northern California and Washington. To test hypotheses related to potential effects from mining land-use activities, salamander larvae were also sampled from a reference site at Whiskeytown National Recreation Area, California, and at a nearby, upstream site (Shasta county) on Bureau of Land Management land where Hg contamination from gold mining activities has been documented. HgT concentrations in whole body larvae ranged from 4.6 to 74.5 ng/g wet wt. and percent MeHg ranged from 67% to 86%. Both HgT and MeHg larval tissue concentrations were significantly higher at the mining site in comparison to measured background levels (P salamander larvae in remote stream ecosystems, where Hg sources were dominated by atmospheric deposition, were generally low in HgT and MeHg and, in comparison, watersheds with a legacy of land-use practices (i.e., mining operations) had approximately 4.5 - 5.5 times the level of HgT bioaccumulation. Moreover, trophic magnification slopes were highest in the Shasta county region where mining was present. These findings suggest that mining activities increase HgT and MeHg exposure to salamander larvae in the region and may present a threat to other higher trophically positioned organisms, and their associated food webs.
Kroll, Andrew J; Garcia, Tiffany S; Jones, Jay E; Dugger, Katie; Murden, Blake; Johnson, Josh; Peterman, Summer; Peerman, Summer; Brintz, Ben; Rochelle, Michael
Plethodontid salamanders are diverse and widely distributed taxa and play critical roles in ecosystem processes. Due to salamander use of structurally complex habitats, and because only a portion of a population is available for sampling, evaluation of sampling designs and estimators is critical to provide strong inference about Plethodontid ecology and responses to conservation and management activities. We conducted a simulation study to evaluate the effectiveness of multi-scale and hierarchical single-scale occupancy models in the context of a Before-After Control-Impact (BACI) experimental design with multiple levels of sampling. Also, we fit the hierarchical single-scale model to empirical data collected for Oregon slender and Ensatina salamanders across two years on 66 forest stands in the Cascade Range, Oregon, USA. All models were fit within a Bayesian framework. Estimator precision in both models improved with increasing numbers of primary and secondary sampling units, underscoring the potential gains accrued when adding secondary sampling units. Both models showed evidence of estimator bias at low detection probabilities and low sample sizes; this problem was particularly acute for the multi-scale model. Our results suggested that sufficient sample sizes at both the primary and secondary sampling levels could ameliorate this issue. Empirical data indicated Oregon slender salamander occupancy was associated strongly with the amount of coarse woody debris (posterior mean = 0.74; SD = 0.24); Ensatina occupancy was not associated with amount of coarse woody debris (posterior mean = -0.01; SD = 0.29). Our simulation results indicate that either model is suitable for use in an experimental study of Plethodontid salamanders provided that sample sizes are sufficiently large. However, hierarchical single-scale and multi-scale models describe different processes and estimate different parameters. As a result, we recommend careful consideration of study questions
Kleinteich, Thomas; Haas, Alexander
Amphibians (Lissamphibia) are characterized by a bi-phasic life-cycle that comprises an aquatic larval stage and metamorphosis to the adult. The ancestral aquatic feeding behavior of amphibian larvae is suction feeding. The negative pressure that is needed for ingestion of prey is created by depression of the hyobranchial apparatus as a result of hyobranchial muscle action. Understanding the homologies of hyobranchial muscles in amphibian larvae is a crucial step in understanding the evolution of this important character complex. However, the literature mostly focuses on the adult musculature and terms used for hyal and ventral branchial muscles in different amphibians often do not reflect homologies across lissamphibian orders. Here we describe the hyal and ventral branchial musculature in larvae of caecilians (Gymnophiona) and salamanders (Caudata), including juveniles of two permanently aquatic salamander species. Based on previous alternative terminology schemes, we propose a terminology for the hyal and ventral branchial muscles that reflects the homologies of muscles and that is suited for studies on hyobranchial muscle evolution in amphibians. We present a discussion of the hyal and ventral branchial muscles in larvae of the most recent common ancestor of amphibians (i.e. the ground plan of Lissamphibia). Based on our terminology, the hyal and ventral branchial musculature of caecilians and salamanders comprises the following muscles: m. depressor mandibulae, m. depressor mandibulae posterior, m. hyomandibularis, m. branchiohyoideus externus, m. interhyoideus, m. interhyoideus posterior, m. subarcualis rectus I, m. subarcualis obliquus II, m. subarcualis obliquus III, m. subarcualis rectus II-IV, and m. transversus ventralis IV. Except for the m. branchiohyoideus externus, all muscles considered herein can be assigned to the ground plan of the Lissamphibia with certainty. The m. branchiohyoideus externus is either apomorphic for the Batrachia (frogs
US Fish and Wildlife Service, Department of the Interior — This outlines the results of field surveys that were conducted for the Cheat Mountain salamander on the Kelley property on three mountains in the Cabin Mountain area...
Michael R. Warburg
Full Text Available This is a long-term study (1974-1999 on the phenology of the rare, xeric- inhabiting salamander Salamandra infraimmaculata in a small isolated population during the breeding season near the breeding ponds on Mt. Carmel. This is a fringe area of the genus’ south-easternmost Palaearctic distribution. Salamanders were captured during the 25 year long study. The first years up to the 1980s the total number of salamanders increased but during the last years there seems to have been a decline. Although this could be a phase in normal population cyclic oscillations nevertheless when compared with long-term data on a European Salamandra it does not seem so. The interpretation of the species’ status is dependent on numbers of salamanders captured as well as on the duration of the study. These subjects are reviewed and discussed in this paper.
Muñoz, David J.; Miller, David A.W.; Sutherland, Chris; Grant, Evan H. Campbell
The cryptic behavior and ecology of herpetofauna make estimating the impacts of environmental change on demography difficult; yet, the ability to measure demographic relationships is essential for elucidating mechanisms leading to the population declines reported for herpetofauna worldwide. Recently developed spatial capture–recapture (SCR) methods are well suited to standard herpetofauna monitoring approaches. Individually identifying animals and their locations allows accurate estimates of population densities and survival. Spatial capture–recapture methods also allow estimation of parameters describing space-use and movement, which generally are expensive or difficult to obtain using other methods. In this paper, we discuss the basic components of SCR models, the available software for conducting analyses, and the experimental designs based on common herpetological survey methods. We then apply SCR models to Red-backed Salamander (Plethodon cinereus), to determine differences in density, survival, dispersal, and space-use between adult male and female salamanders. By highlighting the capabilities of SCR, and its advantages compared to traditional methods, we hope to give herpetologists the resource they need to apply SCR in their own systems.
Geng, Y; Wang, K Y; Zhou, Z Y; Li, C W; Wang, J; He, M; Yin, Z Q; Lai, W M
From February to May 2010, an outbreak of disease occurred amongst farmed Chinese giant salamanders (Andrias davidianus) in Hanzhong County, Shanxi Province, China. Clinical signs included anorexia, lethargy, ecchymoses and swollen areas on the head and limbs, and skin ulceration. The aim of this study was to determine the cause of this disease. Necropsy examination revealed subcutaneous and intramuscular oedema, swollen and pale livers with multifocal haemorrhage, swollen kidneys with multifocal haemorrhage and distended fluid-filled intestines with areas of haemorrhage. Light microscopy revealed intracytoplasmic inclusions suggestive of a viral infection in a variety of organs, as well as degeneration and necrosis of these organs. Electron microscopy of ultrathin sections of the same tissues revealed iridovirus-like particles within the inclusions. Of the six specimens tested, all were positive for ranavirus major capsid protein (MCP) gene. Sequence alignments of the ranavirus MCP gene from these specimens showed 95-98% similarity with published ranavirus data. The virus, provisionally designated as Chinese giant salamander virus (CGSV), was isolated from tissue homogenates of diseased salamanders following inoculation of epithelioma papilloma cyprini cells. Sequence analysis of the MCP genes showed that the isolated virus was a ranavirus with marked sequence identity to other members of the genus Ranavirus. Koch's postulates were fulfilled by infecting healthy Chinese giant salamanders with the CGSV. These salamanders all died within 6-8 days. This is the first report of ranavirus infection associated with mass mortality in Chinese giant salamanders.
Larson, Gary L.; Hoffman, Robert L.
In Mount Rainier National Park, the northwestern salamander usually inhabits relatively large and deep lakes and ponds (average size = 0.3 ha; average depth > 2 m) that contain flocculent, organic bottom sediments and abundant coarse wood. Prior to 1970, salmonids were introduced into many of the park's lakes and ponds that were typical habitat of the northwestern salamander. The objective of this study was to compare, in lakes and ponds with suitable habitat characteristics for northwestern salamanders, the observed abundances of larvae in takes and ponds with and without these introduced salmonids. Day surveys of 61 lakes were conducted between 1993 and 1999. Fish were limited to takes and ponds deeper than 2 in. For the 48 lakes and ponds deeper than 2 in (i.e., 25 fishless lakes and 23 fish lakes), the mean and median observed abundances of northwestern salamander larvae in fishless lakes and ponds was significantly greater than the mean and median observed abundances of larvae in lakes and ponds with fish. Northwestern salamander larvae were not observed in 11 fish lakes. These lakes were similar in median elevation, surface area, and maximum depth to the fishless lakes. The 12 fish lakes with observed larvae were significantly lower in median elevation, larger in median surface area, and deeper in median maximum depth than the fishless lakes. Low to null observed abundances of northwestern salamander larvae in lakes and ponds with fish were attributed to a combination of fish predation of larvae and changes in larval behavior.
Katherine M O'Donnell
Full Text Available Detectability of individual animals is highly variable and nearly always < 1; imperfect detection must be accounted for to reliably estimate population sizes and trends. Hierarchical models can simultaneously estimate abundance and effective detection probability, but there are several different mechanisms that cause variation in detectability. Neglecting temporary emigration can lead to biased population estimates because availability and conditional detection probability are confounded. In this study, we extend previous hierarchical binomial mixture models to account for multiple sources of variation in detectability. The state process of the hierarchical model describes ecological mechanisms that generate spatial and temporal patterns in abundance, while the observation model accounts for the imperfect nature of counting individuals due to temporary emigration and false absences. We illustrate our model's potential advantages, including the allowance of temporary emigration between sampling periods, with a case study of southern red-backed salamanders Plethodon serratus. We fit our model and a standard binomial mixture model to counts of terrestrial salamanders surveyed at 40 sites during 3-5 surveys each spring and fall 2010-2012. Our models generated similar parameter estimates to standard binomial mixture models. Aspect was the best predictor of salamander abundance in our case study; abundance increased as aspect became more northeasterly. Increased time-since-rainfall strongly decreased salamander surface activity (i.e. availability for sampling, while higher amounts of woody cover objects and rocks increased conditional detection probability (i.e. probability of capture, given an animal is exposed to sampling. By explicitly accounting for both components of detectability, we increased congruence between our statistical modeling and our ecological understanding of the system. We stress the importance of choosing survey locations and
Loudon, Andrew H; Holland, Jessica A; Umile, Thomas P; Burzynski, Elizabeth A; Minbiole, Kevin P C; Harris, Reid N
Amphibians possess beneficial skin bacteria that protect against the disease chytridiomycosis by producing secondary metabolites that inhibit the pathogen Batrachochytrium dendrobatidis (Bd). Metabolite production may be a mechanism of competition between bacterial species that results in host protection as a by-product. We expect that some co-cultures of bacterial species or strains will result in greater Bd inhibition than mono-cultures. To test this, we cultured four bacterial isolates (Bacillus sp., Janthinobacterium sp., Pseudomonas sp. and Chitinophaga arvensicola) from red-backed salamanders (Plethodon cinereus) and cultured isolates both alone and together to collect their cell-free supernatants (CFS). We challenged Bd with CFSs from four bacterial species in varying combinations. This resulted in three experimental treatments: (1) CFSs of single isolates; (2) combined CFSs of two isolates; and (3) CFSs from co-cultures. Pair-wise combinations of four bacterial isolates CFSs were assayed against Bd and revealed additive Bd inhibition in 42.2% of trials, synergistic inhibition in 42.2% and no effect in 16.6% of trials. When bacteria isolates were grown in co-cultures, complete Bd inhibition was generally observed, and synergistic inhibition occurred in four out of six trials. A metabolite profile of the most potent co-culture, Bacillus sp. and Chitinophaga arvensicola, was determined with LC-MS and compared with the profiles of each isolate in mono-culture. Emergent metabolites appearing in the co-culture were inhibitory to Bd, and the most potent inhibitor was identified as tryptophol. Thus mono-cultures of bacteria cultured from red-backed salamanders interacted synergistically and additively to inhibit Bd, and such bacteria produced emergent metabolites when cultured together, with even greater pathogen inhibition. Knowledge of how bacterial species interact to inhibit Bd can be used to select probiotics to provide amphibians with protection against Bd.
Andrew Howard Loudon
Full Text Available Amphibians possess beneficial skin bacteria that protect against the disease chytridiomycosis by producing secondary metabolites that inhibit the pathogen Batrachochytrium dendrobatidis (Bd. Metabolite production may be a mechanism of competition between bacterial species that results in host protection as a by-product. We expect that some co-cultures of bacterial species or strains will result in greater Bd inhibition than mono-cultures. To test this, we cultured four bacterial isolates (Bacillus sp., Janthinobacterium sp., Pseudomonas sp. and Chitinophaga arvensicola from red-backed salamanders (Plethodon cinereus and cultured isolates both alone and together to collect their cell-free supernatants (CFS. We challenged Bd with CFSs from four bacterial species in varying combinations. This resulted in three experimental treatments: 1 CFSs of single isolates; 2 combined CFSs of two isolates; and 3 CFSs from co-cultures. Pair-wise combinations of four bacterial isolates CFSs were assayed against Bd and revealed additive Bd inhibition in 42.2% of trials, synergistic inhibition in 42.2% and no effect in 16.6% of trials. When bacteria isolates were grown in co-cultures, complete Bd inhibition was generally observed, and synergistic inhibition occurred in four out of six trials. A metabolite profile of the most potent co-culture, Bacillus sp. and Chitinophaga arvensicola, was determined with LC-MS and compared with the profiles of each isolate in mono-culture. Emergent metabolites appearing in the co-culture were inhibitory to Bd, and the most potent inhibitor was identified as tryptophol. Thus mono-cultures of bacteria cultured from red-backed salamanders interacted synergistically and additively to inhibit Bd, and such bacteria produced emergent metabolites when cultured together, with even greater pathogen inhibition. Knowledge of how bacterial species interact to inhibit Bd can be used to select probiotics to provide amphibians with protection
Ehmcke, Jens; Wistuba, Joachim; Clemen, Günter
The tooth shapes of premaxillary and maxillary teeth of adults of four Mesoamerican salamander species (Urodela, Plethodontidae) were examined. Using scanning electron microscopy we determined whether the monocuspid teeth that are present only on the premaxillary of sexually mature males belong to the conical, unbladed, monocuspid tooth type found in urodele larvae or whether they represent morphological variations of the typical, bladed, bicuspid teeth of metamorphosed individuals. The teeth of some studied species are, in fact, unbladed and in some cases show one very reduced tip. But none of the studied teeth is per definitionem monocuspid and no tooth shows an enameloid layer between dentine shaft and enamel cap, which is always present in the teeth of early urodele larvae. The "monocuspid" teeth of adult males of Mesoamerican plethodontid salamanders have to be considered a morphological variation of the metamorphosed, bicuspid tooth type typical for metamorphosed urodeles.
Skutschas, Pavel P; Baleeva, Nataly V
The development of spinal cord supports (bony thickenings which extend into the vertebral canal of vertebrae) in primitive (Salamandrella keyserlingii) and derived (Lissotriton vulgaris) salamanders were described. The spinal cord supports develop as the protuberances of periostal bone of the neural arches in the anteroproximal part of the septal collagenous fibers which connect a transverse myoseptum with the notochord and spinal cord, in the septal bundle inside the vertebral canal. Spinal cord supports were also found in some teleostean (Salmo salar, Oncorhynchus mykiss) and dipnoan (Protopterus sp.) fishes. The absence of the spinal cord supports in vertebrates with cartilaginous vertebrae (lampreys, chondrichthyan, and chondrostean fishes) corresponds to the fact that the spinal cord supports are bone structures. The absence of the spinal cord supports in frogs correlates with the lack of the well developed septal bundles inside the vertebral canal. The spinal cord supports are, presumably, a synapomorphic character for salamanders which originated independently of those observed in teleostean and dipnoan fishes.
Hossack, Blake R.; Muths, Erin L.; Rorabaugh, James C.; Lemos Espinal, Julio A.; Sigafus, Brent H.; Chambert, Thierry A; Carreon Arroyo, Gerardo; Hurtado Felix, David; Toyos Martinez, Daniel; Jones, Thomas R
The Sonoran Tiger Salamander (Ambystoma mavortium stebbinsi Lowe, 1954) was listed as federally endangered in the USA in 1997 (USFWS 1997). In the USA, the distribution of A. mavortium stebbinsi is limited to the San Rafael Valley (approximately 567 km2), between the Sierra San Antonio (called the Patagonia Mountains in Arizona) and Huachuca Mountains, and south of the Canelo Hills, Arizona (Fig. 1). The USA listing was triggered by loss of natural wetland habitats, threats from invasive predators, frequent die-offs from disease, introgression with the introduced Barred Tiger Salamander (A. mavortium mavortium), and small range and number of breeding sites that increases susceptibility to stochastic events (USFWS 1997). Small population sizes and limited gene flow have caused inbreeding, which may further reduce population viability and the potential for recovery (Jones et al. 1988; Storfer et al. 2014).
A population of facultative neotenous tiger salamanders (A. tigrinum) inhabiting a sewage lagoon at Reese AFB, Hurlwood, Texas, was found to have an exceptionally high rate of spontaneous tissue lesions. The population is composed of an estimated 28,000 large, reproductively mature larvae that are restricted to the lagoon. Only about 17 percent of the population metamorphoses normally. In contrast, tiger salamanders from uncontaminated lagoons in the same general vicinity metamorphose normally; however, no neoplasms were discovered in larvae sampled from the nonsewage lagoons. N-nitrosamine analyses of water and tissue samples of larvae were negative. Polycyclic aromatic hydrocargon analyses revealed traces of benzo(a)pyrene in the sludge; however, perylene, a constituent of jet fuel, was found in high concentration (300 ppB). These results indicate that perylene, which was previously found not be tumorigenic to mice and rats, should be retested as a possible agent for nonmammalian species.
Li WANG; Yong WEI; Yanqing HUANG
Morganella morganii strain 602-1 was isolated from a sick Chinese giant salamander (Andrias davidianus). The strain 602-1 was identiifed through its physiological and biochemical properties and 16S rDNA gene ampliifcation and analyses. Pathogenicity was proven by experimental animal infection, and histopathological examination. The results showed that the ampliifed 16S rDNA sequence of strain 602-1 was 1455 bp, and showed 99%identity with M. morganii. In the host infection experiment, the mortality of Chinese giant salamanders was about 70%. Pathological changes occurred in the spleen, heart, liver, kidney and intestine. This study will help the prevention, understanding and cure for M. morganii infections in amphibians.
Oniscus asellus, 1972-77 . . . . . . 23 6 Mean Oxygen Consumption of the Redbacked Salamander, Plethodon cinereus cinerets, 1972-77 ...... 24 iv During... Plethodon cinereus cinereus (Green), and fotr invertebrates: the earthworm, Lumbricus terrestris L.; the redworm, Lumbricus rubellus (Hoffmeister); the...Mueller. 1972. Diurnal Variation in the Oxygen Consumption of Plethodon cinereus . Amer. Midland Nat., 88: 502-506. Temupleton, J.R. 1960
Zabierek, Kristina C; Gabor, Caitlin R
Prey may use multiple sensory channels to detect predators, whose cues may differ in altered sensory environments, such as turbid conditions. Depending on the environment, prey may use cues in an additive/complementary manner or in a compensatory manner. First, to determine whether the purely aquatic Barton Springs salamander, Eurycea sosorum, show an antipredator response to visual cues, we examined their activity when exposed to either visual cues of a predatory fish (Lepomis cyanellus) or a non-predatory fish (Etheostoma lepidum). Salamanders decreased activity in response to predator visual cues only. Then, we examined the antipredator response of these salamanders to all matched and mismatched combinations of chemical and visual cues of the same predatory and non-predatory fish in clear and low turbidity conditions. Salamanders decreased activity in response to predator chemical cues matched with predator visual cues or mismatched with non-predator visual cues. Salamanders also increased latency to first move to predator chemical cues mismatched with non-predator visual cues. Salamanders decreased activity and increased latency to first move more in clear as opposed to turbid conditions in all treatment combinations. Our results indicate that salamanders under all conditions and treatments preferentially rely on chemical cues to determine antipredator behavior, although visual cues are potentially utilized in conjunction for latency to first move. Our results also have potential conservation implications, as decreased antipredator behavior was seen in turbid conditions. These results reveal complexity of antipredator behavior in response to multiple cues under different environmental conditions, which is especially important when considering endangered species.
Andrew Howard Loudon
Full Text Available Skin bacterial communities can protect amphibians from a fungal pathogen; however, little is known about how these communities are maintained. We used a neutral model of community ecology to identify bacteria that are maintained on salamanders by selection or by dispersal from a bacterial reservoir (soil and ecological drift. We found that 75% (9/12 of bacteria that were consistent with positive selection, < 1% of bacteria that were consistent with random dispersal and none of the bacteria that were consistent under negative selection had a 97% or greater match to antifungal isolates. Additionally we performed an experiment where salamanders were either provided or denied a bacterial reservoir and estimated immigration and loss (emigration and local extinction rates of bacteria on salamanders in both treatments. Loss was strongly related to bacterial richness, suggesting competition is important for structuring the community. Bacteria closely related to antifungal isolates were more likely to persist on salamanders with or without a bacterial reservoir, suggesting they had a competitive advantage. Furthermore, over-represented and under-represented OTUs had similar persistence on salamanders when a bacterial reservoir was present. However, under-represented OTUs were less likely to persist in the absence of a bacterial reservoir, suggesting that the over-represented and under-represented bacteria are selected for or against on salamanders through time. Our findings from the neutral model, migration and persistence analyses show that bacteria that exhibit a high similarity to antifungal isolates persist on salamanders, which likely protect hosts against pathogens and improve fitness. This research is one of the first to apply ecological theory to investigate assembly of host associated-bacterial communities, which can provide insights for probiotic bioaugmentation as a conservation strategy against disease.
Loudon, Andrew H; Venkataraman, Arvind; Van Treuren, William; Woodhams, Douglas C; Parfrey, Laura Wegener; McKenzie, Valerie J; Knight, Rob; Schmidt, Thomas M; Harris, Reid N
Skin bacterial communities can protect amphibians from a fungal pathogen; however, little is known about how these communities are maintained. We used a neutral model of community ecology to identify bacteria that are maintained on salamanders by selection or by dispersal from a bacterial reservoir (soil) and ecological drift. We found that 75% (9/12) of bacteria that were consistent with positive selection, <1% of bacteria that were consistent with random dispersal and none of the bacteria that were consistent under negative selection had a 97% or greater match to antifungal isolates. Additionally we performed an experiment where salamanders were either provided or denied a bacterial reservoir and estimated immigration and loss (emigration and local extinction) rates of bacteria on salamanders in both treatments. Loss was strongly related to bacterial richness, suggesting competition is important for structuring the community. Bacteria closely related to antifungal isolates were more likely to persist on salamanders with or without a bacterial reservoir, suggesting they had a competitive advantage. Furthermore, over-represented and under-represented operational taxonomic units (OTUs) had similar persistence on salamanders when a bacterial reservoir was present. However, under-represented OTUs were less likely to persist in the absence of a bacterial reservoir, suggesting that the over-represented and under-represented bacteria were selected against or for on salamanders through time. Our findings from the neutral model, migration and persistence analyses show that bacteria that exhibit a high similarity to antifungal isolates persist on salamanders, which likely protect hosts against pathogens and improve fitness. This research is one of the first to apply ecological theory to investigate assembly of host associated-bacterial communities, which can provide insights for probiotic bioaugmentation as a conservation strategy against disease.
Marcec, Ruth; Kouba, Andrew; Zhang, Lu; Zhang, Hongxing; Wang, Qijun; Zhao, Hu; Jiang, Wei; Willard, Scott
Worldwide, there are only a handful of reintroduction programs for threatened salamander species, and very few have conducted postrelease studies to examine survival, habitat selection, and dispersal. Limitations in postrelease monitoring are primarily due to size constraints of amphibians and to dimensions of the radiotransmitters available for implantation. However, due to the large size of the critically endangered Chinese giant salamander (Andrias davidianus), these animals make optimal candidates for surgical implantation of radiotransmitters prior to reintroduction or translocation. The objective of this study was to develop an anesthetic protocol using tricane methanesulfonate (MS-222) and test a surgical procedure for coelomic implantation of radiotransmitters for this species. A total of 32 Chinese giant salamanders from two age groups (Group A: 4.7 yr old, n = 16; Group B: 2.7 yr old, n = 16) were implanted with 4-g radiotransmitters designed for underwater monitoring of fish. Group A was held 16 wk before release while Group B was held 6 wk before release, and the salamanders' survival and postoperative complications recorded for the first month postrelease. Group A animals took longer to reach a surgical plane of anesthesia than did Group B animals, and this was directly correlated to mass of the animals. Postsurgery, one animal from Group B died of dehiscence before release while 83.9% animals survived after the first month in the wild. All of the animals that died postrelease were from Group B; three animals experienced dehiscence of the suture site and died while another two animals expired from trauma and fungal infection, respectively. Improvements for future studies include use of alternative suture material for closure after implantation and additional healing time of the incision.
Full Text Available Abstract A re-examination of the mitochondrial genomes of unisexual salamander lineages, published in BMC Evolutionary Biology, shows them to be the oldest unisexual vertebrates known, having been around for 5 million years. This presents a challenge to the prediction that lack of genetic recombination is a fast track to extinction. See research article http://www.biomedcentral.com/1471-2148/10/238
Laura LIMONGI; Gentile Francesco FICETOLA; Giuseppe ROMEO; Raoul MANENTI
Larval growth and survival of organisms are strongly influenced by abiotic and biotic factors, as demonstrated by ex-perimental studies performed under controlled laboratory or semi-natural conditions. Even if they have many advantages, ex-periments cannot cover the full complexity of natural conditions and field studies are needed for a better understanding of how environmental variation determines growth and development rate. Fire salamanderSalamandra salamandrafemales give birth to larvae in a variety of habitats, both epigean and subterranean. In caves, salamander larvae successfully grow and metamorphose, but their growth is more than three times longer than in epigean streams and factors determining these differences require inves-tigation. We performed a field study to understand the factors related to the growth of fire salamander larvae in different envi-ronmental conditions, evaluating the relationship between environmental features and larval growth and differences between caves and epigean spring habitats. Both caves and epigean larvae successfully grew. Capture-mark-recapture allowed to individu-ally track individuals along their whole development, and measure their performance. Growth rate was significantly affected by environmental variables: larvae grew faster in environments with abundant invertebrates and few conspecifics. Taking into ac-count the effect of environmental variables, larval growth was significantly lower in caves. Food availability plays a different ef-fect in the two environments. Larval growth was positively related to the availability of invertebrates in epigean sites only. The development rate of hypogeous populations of salamanders is slower because of multiple parameters, but biotic factors play a much stronger role than the abiotic ones [Current Zoology 61 (3): 421–427, 2015].
Luis Felipe Esqueda
Full Text Available Information available so far is exceedingly meagre about the diet of the snakes included in the genus Liophis, one of the most diverse groups that inhabit terrestrial ecosystems of South America. For the first time is documented the predation of a salamander by Liophis from Venezuela, including a brief overview on the alteration of montane and submontane Andean ecosystem and their effect on the natural dynamic.
Clint R V Otto
Full Text Available Habitat degradation resulting from anthropogenic activities poses immediate and prolonged threats to biodiversity, particularly among declining amphibians. Many studies infer amphibian response to habitat degradation by correlating patterns in species occupancy or abundance with environmental effects, often without regard to the demographic processes underlying these patterns. We evaluated how retention of vertical green trees (CANOPY and coarse woody debris (CWD influenced terrestrial salamander abundance and apparent survival in recently clearcut forests. Estimated abundance of unmarked salamanders was positively related to CANOPY (β Canopy = 0.21 (0.02-1.19; 95% CI, but not CWD (β CWD = 0.11 (-0.13-0.35 within 3,600 m2 sites, whereas estimated abundance of unmarked salamanders was not related to CANOPY (β Canopy = -0.01 (-0.21-0.18 or CWD (β CWD = -0.02 (-0.23-0.19 for 9 m2 enclosures. In contrast, apparent survival of marked salamanders within our enclosures over 1 month was positively influenced by both CANOPY and CWD retention (β Canopy = 0.73 (0.27-1.19; 95% CI and β CWD = 1.01 (0.53-1.50. Our results indicate that environmental correlates to abundance are scale dependent reflecting habitat selection processes and organism movements after a habitat disturbance event. Our study also provides a cautionary example of how scientific inference is conditional on the response variable(s, and scale(s of measure chosen by the investigator, which can have important implications for species conservation and management. Our research highlights the need for joint evaluation of population state variables, such as abundance, and population-level process, such as survival, when assessing anthropogenic impacts on forest biodiversity.
Salvidio, Sebastiano; Palumbi, Giulia; Romano, Antonio; Costa, Andrea
Recent studies suggest that many organisms actively colonize the subterranean environment to avoid climatic stress, exploit new ecological opportunities and reduce competition and predation. Terrestrial salamanders are known to colonize the more stable subterranean habitats mainly to escape external climatic extremes, while the role of predation avoidance remains untested. To better understand the importance of predation, we used clay models of the cave salamander Speleomantes strinatii to compare the predation occurring in woodland and subterranean habitats. Models were positioned in three forests and in three caves in NW Italy. One-hundred eighty-four models were retrieved from the field and 59 (32%) were attacked by predators. Models were attacked on their head more often than expected by chance and, therefore, were perceived by predators as real prey items. In the woodlands, clay models showed a four-time higher probability of being attacked in comparison to caves, suggesting a different level of potential predation risk in these surface habitats. These findings are one of the first experimental evidences that, in terrestrial ecosystems, predation avoidance may contribute to the salamander underground colonization process.
Sutton, William B; Gray, Matthew J; Hoverman, Jason T; Secrist, Richard G; Super, Paul E; Hardman, Rebecca H; Tucker, Jennifer L; Miller, Debra L
Emerging pathogens are a potential contributor to global amphibian declines. Ranaviruses, which infect ectothermic vertebrates and are common in aquatic environments, have been implicated in die-offs of at least 72 amphibian species worldwide. Most studies on the subject have focused on pool-breeding amphibians, and infection trends in other amphibian species assemblages have been understudied. Our primary study objective was to evaluate hypotheses explaining ranavirus prevalence within a lungless salamander assemblage (Family Plethodontidae) in the Great Smoky Mountains National Park, USA. We sampled 566 total plethodontid salamanders representing 14 species at five sites over a 6-year period (2007-2012). We identified ranavirus-positive individuals in 11 of the 14 (78.6%) sampled species, with salamanders in the genus Desmognathus having greatest infection prevalence. Overall, we found the greatest support for site elevation and sampling year determining infection prevalence. We detected the greatest number of infections in 2007 with 82.5% of sampled individuals testing positive for ranavirus, which we attribute to record drought during this year. Infection prevalence remained relatively high in low-elevation sites in 2008 and 2009. Neither body condition nor aquatic dependence was a significant predictor of ranavirus prevalence. Overall, our results indicate that life history differences among species play a minor role determining ranavirus prevalence compared to the larger effects of site elevation and yearly fluctuations (likely due to environmental stressors) during sampling years.
Full Text Available Relationships between species and their habitats are not always constant. Different processes may determine changes in species-habitat association: individuals may prefer different habitat typologies in different periods, or they may be forced to occupy a different habitat in order to follow the changing environment. The aim of our study was to assess whether cave salamanders change their habitat association pattern through the year, and to test whether such changes are determined by environmental changes or by changes in preferences. We monitored multiple caves in Central Italy through one year, and monthly measured biotic and abiotic features of microhabitat and recorded Italian cave salamanders distribution. We used mixed models and niche similarity tests to assess whether species-habitat relationships remain constant through the year. Microhabitat showed strong seasonal variation, with the highest variability in the superficial sectors. Salamanders were associated to relatively cold and humid sectors in summer, but not during winter. Such apparent shift in habitat preferences mostly occurred because the environmental gradient changed through the year, while individuals generally selected similar conditions. Nevertheless, juveniles were more tolerant to dry sectors during late winter, when food demand was highest. This suggests that tolerance for suboptimal abiotic conditions may change through time, depending on the required resources. Differences in habitat use are jointly determined by environmental variation through time, and by changes in the preferred habitat. The trade-offs between tolerance and resources requirement are major determinant of such variation.
Chiari, Ylenia; van der Meijden, Arie; Mucedda, Mauro; Lourenço, João M; Hochkirch, Axel; Veith, Michael
Detecting the factors that determine the interruption of gene flow between populations is key to understanding how speciation occurs. In this context, caves are an excellent system for studying processes of colonization, differentiation and speciation, since they represent discrete geographical units often with known geological histories. Here, we asked whether discontinuous calcareous areas and cave systems represent major barriers to gene flow within and among the five species of Sardinian cave salamanders (genus Hydromantes) and whether intraspecific genetic structure parallels geographic distance within and among caves. We generated mitochondrial cytochrome b gene sequences from 184 individuals representing 48 populations, and used a Bayesian phylogeographic approach to infer possible areas of cladogenesis for these species and reconstruct historical and current dispersal routes among distinct populations. Our results show deep genetic divergence within and among all Sardinian cave salamander species, which can mostly be attributed to the effects of mountains and discontinuities in major calcareous areas and cave systems acting as barriers to gene flow. While these salamander species can also occur outside caves, our results indicate that there is a very poor dispersal of these species between separate cave systems.
Wyderko, Jennie A; Benfield, Ernest F; Maerz, John C; Cecala, Kristen C; Belden, Lisa K
Many factors contribute to parasites varying in host specificity and distribution among potential hosts. Metagonimoides oregonensis is a digenetic trematode that uses stream-dwelling plethodontid salamanders as second intermediate hosts in the Eastern US. We completed a field survey to identify which stream salamander species, at a regional level, are most likely to be important for transmission to raccoon definitive hosts. We surveyed six plethodontid species (N = 289 salamanders) from 23 Appalachian headwater sites in North Carolina: Desmognathus quadramaculatus (n = 69), Eurycea wilderae (n = 160), Desmognathus ocoee (n = 31), Desmognathus monticola (n = 3), Eurycea guttolineata (n = 7), and Gyrinophilus porphyriticus (n = 19). We found infection in all species except D. monticola. Further analysis focused on comparing infection in the two most abundant species, D. quadramaculatus and E. wilderae. We found that D. quadramaculatus had significantly higher infection prevalence and intensity, probably due to a longer aquatic larval period and larger body sizes and thus greater cumulative exposure to the parasite.
Full Text Available Detecting the factors that determine the interruption of gene flow between populations is key to understanding how speciation occurs. In this context, caves are an excellent system for studying processes of colonization, differentiation and speciation, since they represent discrete geographical units often with known geological histories. Here, we asked whether discontinuous calcareous areas and cave systems represent major barriers to gene flow within and among the five species of Sardinian cave salamanders (genus Hydromantes and whether intraspecific genetic structure parallels geographic distance within and among caves. We generated mitochondrial cytochrome b gene sequences from 184 individuals representing 48 populations, and used a Bayesian phylogeographic approach to infer possible areas of cladogenesis for these species and reconstruct historical and current dispersal routes among distinct populations. Our results show deep genetic divergence within and among all Sardinian cave salamander species, which can mostly be attributed to the effects of mountains and discontinuities in major calcareous areas and cave systems acting as barriers to gene flow. While these salamander species can also occur outside caves, our results indicate that there is a very poor dispersal of these species between separate cave systems.
Full Text Available Abstract Background Co-occurrence of distinct colour forms is a classic paradox in evolutionary ecology because both selection and drift tend to remove variation from populations. Apostatic selection, the primary hypothesis for maintenance of colour polymorphism in cryptic animals, proposes that visual predators focus on common forms of prey, resulting in higher survival of rare forms. Empirical tests of this frequency-dependent foraging hypothesis are rare, and the link between predator behaviour and maintenance of variation in prey has been difficult to confirm. Here, we show that predatory birds can act as agents of frequency-dependent selection on terrestrial salamanders. Polymorphism for presence/absence of a dorsal stripe is widespread in many salamander species and its maintenance is a long-standing mystery. Results We used realistic food-bearing model salamanders to test whether selection by wild birds maintains a stripe/no-stripe polymorphism. In experimental manipulations, whichever form was most common was most likely to be attacked by ground-foraging birds, resulting in a survival advantage for the rare form. Conclusion This experiment demonstrates that frequency-dependent foraging by wild birds can maintain colour polymorphism in cryptic prey.
Bogart, J P; Bartoszek, J; Noble, D W A; Bi, K
Although bisexual reproduction has considerable evolutionary benefits, several all-female vertebrates exist. Unisexual salamanders in the genus Ambystoma are common around the Great Lakes region in eastern North America. They originated from a hybridization event that involved a female that shared a common ancestor with Ambystoma barbouri 2.4 to 3.9 million years ago but, unexpectedly, A. barbouri nuclear genomes were unknown in unisexuals. Unisexual salamanders steal sperm from donors of normally bisexual species, so their reproductive mode is described as kleptogenesis. Most known unisexuals are polyploid and they all possess at least one A. laterale genome. One or more other genomes are taken from sperm donors that may include A. jeffersonianum, A. laterale, A. texanum and A. tigrinum. We examined unisexual adults and larvae in a southern Ohio pond where unisexual individuals coexist with male A. barbouri. This population provided an opportunity to test hypotheses pertaining to the role of A. barbouri in the evolution of the disparate cytoplasmic and nuclear genomes in unisexual salamanders. Microsatellite DNA loci, mitochondrial DNA sequences and genomic in situ hybridization were used to identify the genomic constitution of individuals. A. barbouri was found to be an acceptable sperm donor for unisexuals but only contributed genomes in ploidy-elevated individuals. In the absence of A. jeffersonianum, this Ohio population is likely experiencing a recent switch in sperm donors from A. jeffersonianum to A. barbouri and demonstrates the evolutionary flexibility and dynamics of kleptogenesis.
Parish, Clare L; Beljajeva, Anna; Arenas, Ernest; Simon, András
Death and lack of functional regeneration of midbrain dopaminergic (DA) neurons, decreased DA input in the target striatum and movement anomalies characterise Parkinson's disease (PD). There is currently no cure for PD. One way to promote recovery would be to induce or enhance DA neurogenesis. Whether DA neurogenesis occurs in the adult midbrain is a matter of debate. Here, we describe the creation of a salamander 6-hydroxydopamine model of PD to examine midbrain DA regeneration. We demonstrate a robust and complete regeneration of the mesencephalic and diencephalic DA system after elimination of DA neurons. Regeneration is contributed by DA neurogenesis, leads to histological restoration, and to full recovery of motor behaviour. Molecular analyses of the temporal expression pattern of DA determinants indicate that the regenerating DA neurons mature along a similar developmental program as their mammalian counterparts during embryogenesis. We also find that the adult salamander midbrain can reactivate radial glia-like ependymoglia cells that proliferate. The salamander model provides insights into the mechanisms of DA regeneration/neurogenesis and may contribute to the development of novel regenerative strategies for the mammalian brain.
Garza-Garcia, A Acely; Driscoll, Paul C; Brockes, Jeremy P
The most extensive regenerative ability in adult vertebrates is found in the salamanders. Although it is often suggested that regeneration is an ancestral property for vertebrates, our studies on the cell-surface three-finger-protein Prod 1 provide clear evidence for the importance of local evolution of limb regeneration in salamanders. Prod 1 is implicated in both patterning and growth in the regeneration of limbs. It interacts with well-conserved proteins such as the epidermal growth-factor receptor and the anterior gradient protein that are widely expressed in phylogeny. A detailed analysis of the structure and sequence of Prod 1 in relation to other vertebrate three-finger proteins in mammals and zebra fish supports the view that it is a salamander-specific protein. This is the first example of a taxon-specific protein that is clearly implicated in the mechanisms of regeneration. We propose the hypothesis that regeneration depends on the activity of taxon-specific components in orchestrating a cellular machinery that is extensively conserved between regenerating and non-regenerating taxa. This hypothesis has significant implications for our outlook on regeneration in vertebrates, as well as for the strategies employed in extending regenerative ability in mammals.
Bryant Susan V
Full Text Available Abstract Background The basis of genome size variation remains an outstanding question because DNA sequence data are lacking for organisms with large genomes. Sixteen BAC clones from the Mexican axolotl (Ambystoma mexicanum: c-value = 32 × 109 bp were isolated and sequenced to characterize the structure of genic regions. Results Annotation of genes within BACs showed that axolotl introns are on average 10× longer than orthologous vertebrate introns and they are predicted to contain more functional elements, including miRNAs and snoRNAs. Loci were discovered within BACs for two novel EST transcripts that are differentially expressed during spinal cord regeneration and skin metamorphosis. Unexpectedly, a third novel gene was also discovered while manually annotating BACs. Analysis of human-axolotl protein-coding sequences suggests there are 2% more lineage specific genes in the axolotl genome than the human genome, but the great majority (86% of genes between axolotl and human are predicted to be 1:1 orthologs. Considering that axolotl genes are on average 5× larger than human genes, the genic component of the salamander genome is estimated to be incredibly large, approximately 2.8 gigabases! Conclusion This study shows that a large salamander genome has a correspondingly large genic component, primarily because genes have incredibly long introns. These intronic sequences may harbor novel coding and non-coding sequences that regulate biological processes that are unique to salamanders.
Full Text Available The Chinese giant salamander (Andrias davidianus occupies a seat at the phylogenetic and species evolution process, which makes it an invaluable model for genetics; however, the genetic information and gene sequences about the Chinese giant salamander in public databases are scanty. Hence, we aimed to perform transcriptome analysis with the help of high-throughput sequencing. In this data, 61,317,940 raw reads were acquired from Chinese giant salamander mRNA using Illumina paired-end sequencing platform. After de novo assembly, a total of 72,072 unigenes were gained, in which 33,834 (46.95% and 29,479 (40.91% transcripts exhibited homology to sequences in the Nr database and Swiss-Prot database, (E-value <10−5, respectively. In the obtained unigenes, 18,019 (25% transcripts were assigned with at least one Gene Ontology term, of which 1218 (6.8% transcripts were assigned to immune system processes. In addition, a total of 17,572 assembled sequences were assigned into 241 predicted KEGG metabolic pathways. Among these, 2552 (14.5% transcripts were assigned to the immune system relevant pathway and 5 transcripts were identified as potential antimicrobial peptides (AMPs.
Grant, Evan H. Campbell; Muths, Erin L.; Katz, Rachel A.; Canessa, Stefano; Adams, Michael J.; Ballard, Jennifer R.; Berger, Lee; Briggs, Cheryl J.; Coleman, Jeremy; Gray, Matthew J.; Harris, M. Camille; Harris, Reid N.; Hossack, Blake R.; Huyvaert, Kathryn P.; Kolby, Jonathan E.; Lips, Karen R.; Lovich, Robert E.; McCallum, Hamish I.; Mendelson, Joseph R.; Nanjappa, Priya; Olson, Deanna H.; Powers, Jenny G.; Richgels, Katherine L.D.; Russell, Robin E.; Schmidt, Benedikt R.; Spitzen-van der Sluijs, Annemarieka; Watry, Mary Kay; Woodhams, Douglas C.; White, C. LeAnn
The recently (2013) identified pathogenic chytrid fungus, Batrachochytrium salamandrivorans (Bsal), poses a severe threat to the distribution and abundance of salamanders within the United States and Europe. Development of a response strategy for the potential, and likely, invasion of Bsal into the United States is crucial to protect global salamander biodiversity. A formal working group, led by Amphibian Research and Monitoring Initiative (ARMI) scientists from the U.S. Geological Survey (USGS) Patuxent Wildlife Research Center, Fort Collins Science Center, and Forest and Rangeland Ecosystem Science Center, was held at the USGS Powell Center for Analysis and Synthesis in Fort Collins, Colorado, United States from June 23 to June 25, 2015, to identify crucial Bsal research and monitoring needs that could inform conservation and management strategies for salamanders in the United States. Key findings of the workshop included the following: (1) the introduction of Bsal into the United States is highly probable, if not inevitable, thus requiring development of immediate short-term and long-term intervention strategies to prevent Bsal establishment and biodiversity decline; (2) management actions targeted towards pathogen containment may be ineffective in reducing the long-term spread of Bsal throughout the United States; and (3) early detection of Bsal through surveillance at key amphibian import locations, among high-risk wild populations, and through analysis of archived samples is necessary for developing management responses. Top research priorities during the preinvasion stage included the following: (1) deployment of qualified diagnostic methods for Bsal and establishment of standardized laboratory practices, (2) assessment of susceptibility for amphibian hosts (including anurans), and (3) development and evaluation of short- and long-term pathogen intervention and management strategies. Several outcomes were achieved during the workshop, including development
Sato, Hiroshi; Ihara, Sadao; Inaba, Osamu; Une, Yumi
Since 2000, cutaneous nodular lesions were observed in Tohoku hynobiid salamanders (Hynobius lichenatus) in the northern Abukuma Mountains, northeastern Honshu, Japan. The distribution of diseased salamanders has expanded, along with the increasing severity of the disease. Samples (foreleg, hindleg, and tail) from five diseased salamanders caught in the spring of 2009 contained 11 to 383 metacercarial cysts (0.43-0.48 mm in diameter) per individual. Experimental inoculations of the metacercariae into rats (Rattus norvegicus) resulted in the development of pentagonal adults with intrauterine eggs that were collected at 3 days, 2 wk, and 5 wk postinoculation. Although adults obtained from rats were smaller, the flukes were morphologically similar to Euryhelmis costaricensis, recorded from Japanese martens (Martes melampus) in northeastern Honshu. The ribosomal RNA gene sequences of the metacercariae collected from salamanders and the E. costaricensis collected from a Japanese marten were identical, confirming the morphology-based species identification. Environmental changes responsible for the increased frequency of diseased salamanders and the increased parasite load are unknown, although increases in feral populations of introduced raccoons (Procyon lotor) and American minks (Neovison vison) may be associated.
Hossack, Blake R.; Honeycutt, Richard; Sigafus, Brent H.; Muths, Erin L.; Crawford, Catherine L.; Jones, Thomas R.; Sorensen, Jeff A.; Rorabaugh, James C.; Chambert, Thierry
Understanding the additive or interactive threats of habitat transformation and invasive species is critical for conservation, especially where climate change is expected to increase the severity or frequency of drought. In the arid southwestern USA, this combination of stressors has caused widespread declines of native aquatic and semi-aquatic species. Achieving resilience to drought and other effects of climate change may depend upon continued management, so understanding the combined effects of stressors is important. We used Bayesian hierarchical models fitted with 10-years of pond-based monitoring surveys for the federally-endangered Sonoran Tiger Salamander (Ambystoma mavortium stebbinsi) and invasive predators (fishes and American Bullfrogs, Lithobates catesbeianus) that threaten native species. We estimated trends in occupancy of salamanders and invasive predators while accounting for hydrological dynamics of ponds, then used a two-species interaction model to directly estimate how invasive predators affected salamander occupancy. We also tested a conceptual model that predicted that drought, by limiting the distribution of invasive predators, could ultimately benefit native species. Even though occupancy of invasive predators was stationary and their presence in a pond reduced the probability of salamander presence by 23%, occupancy of Sonoran Tiger Salamanders increased, annually, by 2.2%. Occupancy of salamanders and invasive predators both declined dramatically following the 5th consecutive year of drought. Salamander occupancy recovered quickly after return to non-drought conditions, while occupancy of invasive predators remained suppressed. Models that incorporated three time-lagged periods (1 to 4 years) of local moisture conditions confirmed that salamanders and invasive predators responded differently to drought, reflecting how life-history strategies shape responses to disturbances. The positive 10-year trend in salamander occupancy and their
Huston, D C; Cantu, V; Huffman, D G
The gill parasite Centrocestus formosanus (Trematoda: Heterophyidae) is an exotic parasite of concern in Texas because it has been shown to infect multiple threatened and endangered fish species. The purpose of this study was to determine if C. formosanus could present a threat to larval anurans, as well as threatened neotenic salamanders endemic to the spring-fed systems of Texas. We exposed adults of the San Marcos salamander Eurycea nana (Caudata: Plethodontidae) and tadpoles of the Rio Grande leopard frog Lithobates berlandieri (Anura: Ranidae) to the cercariae of C. formosanus . The San Marcos salamander showed no signs of metacercarial infection, suggesting that E. nana may be refractory to C. formosanus cercariae. Centrocestus formosanus readily infects the gills of leopard frog tadpoles, but the metacercariae apparently died prior to reaching maturity in our tadpoles.
Ju, B G; Kim, W S
Cathepsin D is a major lysosomal aspartic proteinase participating in the degradation or modification of intra- and extracellular matrix molecules, and its activity is known to increase in the process of tissue reorganization during the early phase of salamander limb regeneration. Here, we report the cloning of a salamander cathepsin D cDNA from Hynobius leechii and its expression profile in normal and retinoic acid (RA) treated limb regenerates. The gene expression of cathepsin D increased notably during the dedifferentiation stage and decreased gradually thereafter. Furthermore, RA that enhances dedifferentiation of regenerating salamander limb caused the elevation of cathepsin D expression both in terms of level and duration. These results suggest that cathepsin D plays important role(s) in the dedifferentiation process, and enhancement of cathepsin D expression might be closely related to RA-evoked pattern duplication.
Full Text Available Worldwide, one in every three species of amphibian is endangered, 39 species have gone extinct in the last 500 years and another 130 species are suspected to have gone extinct in recent decades. Of the amphibians, salamanders have the highest portion of their species in one of the risk categories, even higher than the frogs. To date there have been few studies that have used recent field data to examine the status of populations of endangered salamanders. In this study we evaluate the current situation of two tiny salamanders, Parvimolge townsendi and Thorius pennatulus, both of which are distributed at intermediate elevations in the mountains of the northern Neotropics and are considered to be critically endangered; the first has been proposed as possibly extinct. By carrying out exhaustive surveys in both historical and potentially suitable sites for these two species, we evaluated their abundance and the characteristics of their habitats, and we estimated their potential geographic distribution. We visited 22 sites, investing 672 person-hours of sampling effort in the surveys, and found 201 P. townsendi salamanders in 11 sites and only 13 T. pennatulus salamanders in 5 sites. Both species were preferentially found in cloud forest fragments that were well conserved or only moderately transformed, and some of the salamanders were found in shade coffee plantations. The potential distribution area of both species is markedly fragmented and we estimate that it has decreased by more than 48%. The results of this study highlight the importance of carrying out exhaustive, systematic field surveys to obtain accurate information about the current situation of critically endangered species, and help us better understand the crisis that amphibians are facing worldwide.
Kenison, Erin K; Litt, Andrea R.; Pilliod, David; McMahon, Tom E
Predation by nonnative fishes has reduced abundance and increased extinction risk for amphibian populations worldwide. Although rare, fish and palatable amphibians have been observed to coexist where aquatic vegetation and structural complexity provide suitable refugia. We examined whether larval long-toed salamanders (Ambystoma macrodactylum Baird, 1849) increased use of vegetation cover in lakes with trout and whether adding vegetation structure could reduce predation risk and nonconsumptive effects (NCEs), such as reductions in body size and delayed metamorphosis. We compared use of vegetation cover by larval salamanders in lakes with and without trout and conducted a field experiment to investigate the influence of added vegetation structure on salamander body morphology and life history. The probability of catching salamanders in traps in lakes with trout was positively correlated with the proportion of submerged vegetation and surface cover. Growth rates of salamanders in enclosures with trout cues decreased as much as 85% and the probability of metamorphosis decreased by 56%. We did not find evidence that adding vegetation reduced NCEs in experimental enclosures, but salamanders in lakes with trout utilized more highly-vegetated areas which suggests that adding vegetation structure at the scale of the whole lake may facilitate coexistence between salamanders and introduced trout.
Courtney L. Davis,; David A.W. Miller,; Walls, Susan; Barichivich, William J.; Riley, Jeffrey W.; Brown, Mary E.
Plasticity in life history strategies can be advantageous for species that occupy spatially or temporally variable environments. We examined how phenotypic plasticity influences responses of the mole salamander, Ambystoma talpoideum, to disturbance events at the St. Marks National Wildlife Refuge (SMNWR), FL, USA from 2009 to 2014. We observed periods of extensive drought early in the study, in contrast to high rainfall and expansive flooding events in later years. Flooding facilitated colonization of predatory fishes to isolated wetlands across the refuge. We employed multistate occupancy models to determine how this natural experiment influenced the occurrence of aquatic larvae and paedomorphic adults and what implications this may have for the population. We found that, in terms of occurrence, responses to environmental variation differed between larvae and paedomorphs, but plasticity (i.e. the ability to metamorphose rather than remain in aquatic environment) was not sufficient to buffer populations from declining as a result of environmental perturbations. Drought and fish presence negatively influenced occurrence dynamics of larval and paedomorphic mole salamanders and, consequently, contributed to observed short-term declines of this species. Overall occurrence of larval salamanders decreased from 0.611 in 2009 to 0.075 in 2014 and paedomorph occurrence decreased from 0.311 in 2009 to 0.121 in 2014. Although variation in selection pressures has likely maintained this polyphenism previously, our results suggest that continued changes in environmental variability and the persistence of fish in isolated wetlands could lead to a loss of paedomorphosis in the SMNWR population and, ultimately, impact regional persistence in the future.
Davis, Drew R; Gabor, Caitlin R
Exposure to predatory stimuli typically results in the elevation of circulating glucocorticoid levels and a behavioral response of freezing or escape behavior in many prey species. Corticosterone (CORT) is the main glucocorticoid in amphibians and is known to be important in modulating many behaviors and developmental functions. The federally threatened San Marcos salamander, Eurycea nana, decreases activity in response to both native and introduced predatory fish, however, experience may further influence these interactions. To better understand the indirect effects of fish predators on this salamander, we examined both the antipredator behavior and water-borne CORT release rates in response to chemical cues (kairomones) from two fish species that varied in temporal risk of predation: (1) a low encounter frequency predator (largemouth bass, Micropterus salmoides), (2) a high encounter frequency predator (redbreast sunfish, Lepomis auritus), and (3) a blank water control. Salamanders reduced activity (antipredator response) after exposure to both predator treatments, but not to the blank water control, and the response to M. salmoides was significantly stronger than that to L. auritus. The CORT response (post-stimulus/pre-stimulus release rates) did not differ between the blank water control and L. auritus treatments, and both were significantly less than the CORT response to M. salmoides. Overall, E. nana showed a decreased antipredator response and no CORT response towards the high encounter frequency L. auritus as compared to the low encounter frequency M. salmoides. Eurycea nana may mute antipredator and CORT responses to high temporal frequency predators. There was, however, no correlation between CORT release rates and antipredator behavior, which suggests that the presence of predators may be affecting CORT response and behavior independently.
Pilliod, David S.; Arkle, Robert S.; Maxell, Bryce A.
Studies have demonstrated negative effects of non-native, predatory fishes on native amphibians, yet it is still unclear why some amphibian populations persist, while others are extirpated, following fish invasion. We examined this question by developing habitat-based occupancy models for the long-toed salamander (Ambystoma macrodactylum) and nonnative fish using survey data from 1,749 water bodies across 470 catchments in the Northern Rocky Mountains, USA. We first modeled the habitat associations of salamanders at 468 fishless water bodies in 154 catchments where non-native fish were historically, and are currently, absent from the entire catchment. Wethen applied this habitat model to the complete data set to predict the probability of salamander occupancy in each water body, removing any effect of fish presence. Finally, we compared field-observed occurrences of salamanders and fish to modeled probability of salamander occupancy. Suitability models indicated that fish and salamanders had similar habitat preferences, possibly resulting in extirpations of salamander populations from entire catchments where suitable habitats were limiting. Salamanders coexisted with non-native fish in some catchments by using marginal quality, isolated (no inlet or outlet) habitats that remained fishless. They rarely coexisted with fish within individual water bodies and only where habitat quality was highest. Connectivity of water bodies via streams resulted in increased probability of fish invasion and consequently reduced probability of salamander occupancy.These results could be used to identify and prioritize catchments and water bodies where control measures would be most effective at restoring amphibian populations. Our approach could be useful as a framework for improved investigations into questions of persistence and extirpation of native species when non-native species have already become established.
Reinhardt, Timm; Steinfartz, Sebastian; Paetzold, Achim; Weitere, Markus
Shifts in life history traits and in the behaviour of species can potentially alter ecosystem functioning. The reproduction of the central European fire salamander (Salamandra salamandra), which usually deposits its larvae in first-order streams, in small pool and pond-like habitats, is an example of a recent local adaptation in this species. Here we aimed to quantify the direct and indirect effects of the predatory larvae on the aquatic food webs in the ponds and on the flux of matter between the ponds and adjacent terrestrial habitats. Our estimates are based on biomass data of the present pond fauna as well as on the analysis of stomach content data, growth rates and population dynamics of the salamander larvae in pond habitats. By their deposition of larvae in early spring, female fire salamanders import between 0.07 and 2.86 g dry mass m(-2) larval biomass into the ponds. Due to high mortality rates in the larval phase and the relatively small size at metamorphosis of the pond-adapted salamanders compared to stream-adapted ones, the biomass export of the metamorphosed salamanders clearly falls below the initial biomass import. Catastrophic events such as high water temperatures and low oxygen levels may even occasionally result in mass mortalities of salamander larvae and thus in a net 100 % import of the salamander biomass into the pond food webs. Indirect effects further accelerate this net import of matter into the aquatic habitat, e.g. the feeding of salamanders on aquatic insect larvae with the emergence of terrestrial adults-thus preventing export-and on terrestrial organisms that fall on the water surface (supporting import). This study demonstrates that the adaptation of salamanders to pond reproduction can alter food web linkages across ecosystem boundaries by enhancing the flux of materials and energy from terrestrial (i.e. forest) to the aquatic (i.e. pond) habitat.
Caruso, Nicholas M; Sears, Michael W; Adams, Dean C; Lips, Karen R
Reduction in body size is a major response to climate change, yet evidence in globally imperiled amphibians is lacking. Shifts in average population body size could indicate either plasticity in the growth response to changing climates through changes in allocation and energetics, or through selection for decreased size where energy is limiting. We compared historic and contemporary size measurements in 15 Plethodon species from 102 populations (9450 individuals) and found that six species exhibited significant reductions in body size over 55 years. Biophysical models, accounting for actual changes in moisture and air temperature over that period, showed a 7.1-7.9% increase in metabolic expenditure at three latitudes but showed no change in annual duration of activity. Reduced size was greatest at southern latitudes in regions experiencing the greatest drying and warming. Our results are consistent with a plastic response of body size to climate change through reductions in body size as mediated through increased metabolism. These rapid reductions in body size over the past few decades have significance for the susceptibility of amphibians to environmental change, and relevance for whether adaptation can keep pace with climate change in the future.
Arntzen, J.W.; Beukema, W.; Galis, F.; Ivanović, A.
In vertebrates, the relative proportion of the number of trunk and caudal vertebrae is an important determinant of body shape. While among amphibians frogs and toads show low variation in vertebrae numbers, in salamanders the numbers of trunk and caudal vertebrae vary widely, giving rise to phenotyp
Full Text Available Urbanization is a major cause of amphibian decline. Stream-dwelling plethodontid salamanders are particularly susceptible to urbanization due to declining water quality and hydrological changes, but few studies have examined these taxa in cities. The northern dusky salamander (Desmognathus fuscus was once common in the New York City metropolitan area, but has substantially declined throughout the region in recent decades. We used five tetranucleotide microsatellite loci to examine population differentiation, genetic variation, and bottlenecks among five remnant urban populations of dusky salamanders in NYC. These genetic measures provide information on isolation, prevalence of inbreeding, long-term prospects for population persistence, and potential for evolutionary responses to future environmental change. All populations were genetically differentiated from each other, and the most isolated populations in Manhattan have maintained very little genetic variation (i.e. <20% heterozygosity. A majority of the populations also exhibited evidence of genetic bottlenecks. These findings contrast with published estimates of high genetic variation within and lack of structure between populations of other desmognathine salamanders sampled over similar or larger spatial scales. Declines in genetic variation likely resulted from population extirpations and the degradation of stream and terrestrial paths for dispersal in NYC. Loss of genetic variability in populations isolated by human development may be an underappreciated cause and/or consequence of the decline of this species in urbanized areas of the northeast USA.
Matsui, Masafumi; Tominaga, Atsushi; Liu, Wan-zhao; Tanaka-Ueno, Tomoko
The phylogenetic relationships among 46 samples from 27 populations of the Japanese giant salamander, Andriasjaponicus and its congener, A. davidianus from China was investigated, using 3664 bp sequences of the mitochondrial genes NADH1, NADH3, cyt b and CR, partial NADH6 and intervening genes. In phylogenetic trees constructed by MP, ML, and Bayesian methods, the family Cryptobranchidae and the genus Andrias both form monophyletic groups. Japanese A. japonicus and Chinese A. davidianus are sister taxa and can be regarded as separate species despite a small degree of genetic differentiation. Andriasjaponicus is divided into central and western clades, but the phylogenetic relationships within the latter clade are unresolved. As previously reported from allozyme analyses, A. japonicus exhibits little genetic differentiation, in strong contrast to salamanders of the genus Hynobius with which their distributions overlap. This reduced genetic variability in A. japonicus is attributable to a unique mating system of polygyny, delayed sexual maturity, notable longevity, life in a stable aquatic environment, and gigantism, as well as bottleneck effects following habitat fragmentation and extinction of local populations during Quaternary glaciations. The species is thus susceptible to extinction by potential environmental fluctuations, and requires extensive conservation measures.
Lamb, Jennifer Y.; Waddle, J. Hardin; Qualls, Carl P.
Large gaps exist in our knowledge of the ecology of stream-breeding plethodontid salamanders in the Gulf Coastal Plain. Data describing where these salamanders are likely to occur along environmental gradients, as well as their likelihood of detection, are important for the prevention and management of amphibian declines. We used presence/absence data from leaf litter bag surveys and a hierarchical Bayesian multispecies single-season occupancy model to estimate the occurrence of five species of plethodontids across reaches in headwater streams in the Gulf Coastal Plain. Average detection probabilities were high (range = 0.432–0.942) and unaffected by sampling covariates specific to the use of litter bags (i.e., bag submergence, sampling season, in-stream cover). Estimates of occurrence probabilities differed substantially between species (range = 0.092–0.703) and were influenced by the size of the upstream drainage area and by the maximum proportion of the reach that dried. The effects of these two factors were not equivalent across species. Our results demonstrate that hierarchical multispecies models successfully estimate occurrence parameters for both rare and common stream-breeding plethodontids. The resulting models clarify how species are distributed within stream networks, and they provide baseline values that will be useful in evaluating the conservation statuses of plethodontid species within lotic systems in the Gulf Coastal Plain.
David B. Wake
Full Text Available Here we characterize the genetic structure of Black Salamanders (Aneides flavipunctatus in the Klamath Mountains of northwestern California and southwestern Oregon using mitochondrial and nuclear DNA sequences. We hypothesized that the Sacramento, Smith, Klamath, and Rogue River watersheds would represent distinct genetic populations based on prior ecological results, which suggest that Black Salamanders avoid high elevations such as the ridges that separate watersheds. Our mitochondrial results revealed two major lineages, one in the Sacramento River watershed, and another containing the Klamath, Smith, and Rogue River watersheds. Clustering analyses of our thirteen nuclear loci show the Sacramento watershed population to be genetically distinctive. Populations in the Klamath, Smith, and Rogue watersheds are also distinctive but not as differentiated and their boundaries do not correspond to watersheds. Our historical demographic analyses suggest that the Sacramento population has been isolated from the Klamath populations since the mid-Pleistocene, with negligible subsequent gene flow (2 Nm ≤ 0.1. The Smith and Rogue River watershed populations show genetic signals of recent population expansion. These results suggest that the Sacramento River and Klamath River watersheds served as Pleistocene refugia, and that the Rogue and Smith River watersheds were colonized more recently by northward range expansion from the Klamath.
Blooi, M; Pasmans, F; Rouffaer, L; Haesebrouck, F; Vercammen, F; Martel, A
Chytridiomycosis caused by the chytrid fungus Batrachochytrium salamandrivorans (Bsal) poses a serious threat to urodelan diversity worldwide. Antimycotic treatment of this disease using protocols developed for the related fungus Batrachochytrium dendrobatidis (Bd), results in therapeutic failure. Here, we reveal that this therapeutic failure is partly due to different minimum inhibitory concentrations (MICs) of antimycotics against Bsal and Bd. In vitro growth inhibition of Bsal occurs after exposure to voriconazole, polymyxin E, itraconazole and terbinafine but not to florfenicol. Synergistic effects between polymyxin E and voriconazole or itraconazole significantly decreased the combined MICs necessary to inhibit Bsal growth. Topical treatment of infected fire salamanders (Salamandra salamandra), with voriconazole or itraconazole alone (12.5 μg/ml and 0.6 μg/ml respectively) or in combination with polymyxin E (2000 IU/ml) at an ambient temperature of 15 °C during 10 days decreased fungal loads but did not clear Bsal infections. However, topical treatment of Bsal infected animals with a combination of polymyxin E (2000 IU/ml) and voriconazole (12.5 μg/ml) at an ambient temperature of 20 °C resulted in clearance of Bsal infections. This treatment protocol was validated in 12 fire salamanders infected with Bsal during a field outbreak and resulted in clearance of infection in all animals.
Ousterhout, Brittany H; Semlitsch, Raymond D
Conditions experienced in early developmental stages can have long-term consequences for individual fitness. High intraspecific density during the natal period can affect juvenile and eventually adult growth rates, metabolism, immune function, survival, and fecundity. Despite the important ecological and evolutionary effects of early developmental density, the form of the relationship between natal density and resulting juvenile phenotype is poorly understood. To test competing hypotheses explaining responses to intraspecific density, we experimentally manipulated the initial larval density of ringed salamanders (Ambystoma annulatum), a pond-breeding amphibian, over 11 densities. We modeled the functional form of the relationship between natal density and juvenile traits, and compared the relative support for the various hypotheses based on their goodness of fit. These functional form models were then used to parameterize a simple simulation model of population growth. Our data support non-additive density dependence and presents an alternate hypothesis to additive density dependence, self-thinning and Allee effects in larval amphibians. We posit that ringed salamander larvae may be under selective pressure for tolerance to high density and increased efficiency in resource utilization. Additionally, we demonstrate that models of population dynamics are sensitive to assumptions of the functional form of density dependence.
Bonett, Ronald M; Steffen, Michael A; Lambert, Shea M; Wiens, John J; Chippindale, Paul T
Life-history modes can profoundly impact the biology of a species, and a classic example is the dichotomy between metamorphic (biphasic) and paedomorphic (permanently aquatic) life-history strategies in salamanders. However, despite centuries of research on this system, several basic questions about the evolution of paedomorphosis in salamanders have not been addressed. Here, we use a nearly comprehensive, time-calibrated phylogeny of spelerpine plethodontids to reconstruct the evolution of paedomorphosis and to test if paedomorphosis is (1) reversible; (2) associated with living in caves; (3) associated with relatively dry climatic conditions on the surface; and (4) correlated with limited range size and geographic dispersal. We find that paedomorphosis arose multiple times in spelerpines. We also find evidence for re-evolution of metamorphosis after several million years of paedomorphosis in a lineage of Eurycea from the Edwards Plateau region of Texas. We also show for the first time using phylogenetic comparative methods that paedomorphosis is highly correlated with cave-dwelling, arid surface environments, and small geographic range sizes, providing insights into both the causes and consequences of this major life history transition.
Nathan F Bendik
Full Text Available Despite recognition that nearly one-third of the 6300 amphibian species are threatened with extinction, our understanding of the general ecology and population status of many amphibians is relatively poor. A widely-used method for monitoring amphibians involves injecting captured individuals with unique combinations of colored visible implant elastomer (VIE. We compared VIE identification to a less-invasive method - computer-assisted photographic identification (photoID - in endangered Jollyville Plateau salamanders (Eurycea tonkawae, a species with a known range limited to eight stream drainages in central Texas. We based photoID on the unique pigmentation patterns on the dorsal head region of 1215 individual salamanders using identification software Wild-ID. We compared the performance of photoID methods to VIEs using both 'high-quality' and 'low-quality' images, which were taken using two different camera types and technologies. For high-quality images, the photoID method had a false rejection rate of 0.76% compared to 1.90% for VIEs. Using a comparable dataset of lower-quality images, the false rejection rate was much higher (15.9%. Photo matching scores were negatively correlated with time between captures, suggesting that evolving natural marks could increase misidentification rates in longer term capture-recapture studies. Our study demonstrates the utility of large-scale capture-recapture using photo identification methods for Eurycea and other species with stable natural marks that can be reliably photographed.
Marshall, Jeremy L.; Camp, Carlos D.
Biological diversity is distributed across the planet in non-random, organised ways. At the species level, numerous environmental variables have been proposed to explain this non-random distribution with available energy and habitat heterogeneity receiving the most empirical support. With regard to genetic organisation, environmental stress and habitat heterogeneity have been widely supported. However, few studies have addressed if these two scales of biological organisation are structured via similar processes. Here, we tested whether or not the distributional organisation of genetic and species richness were driven by similar environmental variables for salamanders of the family Plethodontidae across North America. In general, we found that those environmental variables related to energy, particularly energy made accessible to salamanders via the actions of available moisture, were the primary determinants of both genetic and species richness. This finding is consistent with both the "more individuals hypothesis" of species richness and neutral-theory expectations for genetic richness. Additionally, greater habitat heterogeneity, as measured by increased topographic variance, was of secondary importance in positively influencing species richness, although its effects on genetic richness were far more variable. In total, our results suggest that both of these scales of biological organisation are influenced by similar environmental variables, even though increased genetic richness at the population-level does not always translate into greater species richness.
Lu, Bin; Zheng, Yuchi; Murphy, Robert W; Zeng, Xiaomao
Orogenesis of topographically diverse montane regions often drives complex evolutionary histories of species. The extensive biodiversity of the eastern edge of the Tibetan Plateau, which gradually decreases eastwardly, facilitates a comparison of historical patterns. We use coalescence methods to compare species of stream salamanders (Batrachuperus) that occur at high and low elevations. Coalescent simulations reveal that closely related species are likely to have been influenced by different drivers of diversification. Species living in the western high-elevation region with its northsouth extending mountains appear to have experienced colonization via dispersal followed by isolation and divergence. In contrast, species on the eastern low-elevation region, which has many discontinuous mountain ranges, appear to have experienced fragmentation, sometimes staged, of wide-ranging ancestral populations. The two groups of species appear to have been affected differently by glaciation. High-elevation species, which are more resistant to cooler temperatures, appear to have experienced population declines as recently as the last glaciation (0.016-0.032Ma). In contrast, salamanders dwelling in the warmer and wetter habitats at low-elevation environs appear to have been affected less by the relatively recent, milder glaciation, and more so by harsher, extensive glaciations (0.5-0.175 Ma). Thus, elevation, topography and cold tolerance appear to drive evolutionary patterns of diversification and demography even among closely related taxa. The comparison of multiple species in genealogical analyses can lead to an understanding of the evolutionary drivers.
O'Donnell, Katherine; Messerman, Arianne F; Barichivich, William J.; Semlitsch, Raymond D.; Gorman, Thomas A.; Mitchell, Harold G; Allan, Nathan; Fenolio, Dante B.; Green, Adam; Johnson, Fred A.; Keever, Allison; Mandica, Mark; Martin, Julien; Mott, Jana; Peacock, Terry; Reinman, Joseph; Romanach, Stephanie; Titus, Greg; McGowan, Conor; Walls, Susan
At least one-third of all amphibian species face the threat of extinction, and current amphibian extinction rates are four orders of magnitude greater than background rates. Preventing extirpation often requires both ex situ (i.e., conservation breeding programs) and in situ strategies (i.e., protecting natural habitats). Flatwoods salamanders (Ambystoma bishopi and A. cingulatum) are protected under the U.S. Endangered Species Act. The two species have decreased from 476 historical locations to 63 recently extant locations (86.8% loss). We suggest that recovery efforts are needed to increase populations and prevent extinction, but uncertainty regarding optimal actions in both ex situ and in situ realms hinders recovery planning. We used structured decision making (SDM) to address key uncertainties regarding both captive breeding and habitat restoration, and we developed short-, medium-, and long-term goals to achieve recovery objectives. By promoting a transparent, logical approach, SDM has proven vital to recovery plan development for flatwoods salamanders. The SDM approach has clear advantages over other previous approaches to recovery efforts, and we suggest that it should be considered for other complex decisions regarding endangered species.
Zhu, Wei; Kuo, Dwight; Nathanson, Jason; Satoh, Akira; Pao, Gerald M; Yeo, Gene W; Bryant, Susan V; Voss, S Randal; Gardiner, David M; Hunter, Tony
Salamanders possess an extraordinary capacity for tissue and organ regeneration when compared to mammals. In our effort to characterize the unique transcriptional fingerprint emerging during the early phase of salamander limb regeneration, we identified transcriptional activation of some germline-specific genes within the Mexican axolotl (Ambystoma mexicanum) that is indicative of cellular reprogramming of differentiated cells into a germline-like state. In this work, we focus on one of these genes, the long interspersed nucleotide element-1 (LINE-1) retrotransposon, which is usually active in germ cells and silent in most of the somatic tissues in other organisms. LINE-1 was found to be dramatically upregulated during regeneration. In addition, higher genomic LINE-1 content was also detected in the limb regenerate when compared to that before amputation indicating that LINE-1 retrotransposition is indeed active during regeneration. Active LINE-1 retrotransposition has been suggested to have a potentially deleterious impact on genomic integrity. Silencing of activated LINE-1 by small RNAs has been reported to be part of the machinery aiming to maintain genomic integrity. Indeed, we were able to identify putative LINE-1-related piRNAs in the limb blastema. Transposable element-related piRNAs have been identified frequently in the germline in other organisms. Thus, we present here a scenario in which a unique germline-like state is established during axolotl limb regeneration, and the re-activation of LINE-1 may serve as a marker for cellular dedifferentiation in the early-stage of limb regeneration.
Monaghan, James R; Walker, John A; Page, Robert B; Putta, Srikrishna; Beachy, Christopher K; Voss, S Randal
In contrast to mammals, salamanders have a remarkable ability to regenerate their spinal cord and recover full movement and function after tail amputation. To identify genes that may be associated with this greater regenerative ability, we designed an oligonucleotide microarray and profiled early gene expression during natural spinal cord regeneration in Ambystoma mexicanum. We sampled tissue at five early time points after tail amputation and identified genes that registered significant changes in mRNA abundance during the first 7 days of regeneration. A list of 1036 statistically significant genes was identified. Additional statistical and fold change criteria were applied to identify a smaller list of 360 genes that were used to describe predominant expression patterns and gene functions. Our results show that a diverse injury response is activated in concert with extracellular matrix remodeling mechanisms during the early acute phase of natural spinal cord regeneration. We also report gene expression similarities and differences between our study and studies that have profiled gene expression after spinal cord injury in rat. Our study illustrates the utility of a salamander model for identifying genes and gene functions that may enhance regenerative ability in mammals.
Um, JungIn; Jung, Da-Woon; Williams, Darren Reece
The ability of salamanders, such as newts, to regenerate damaged tissues has been studied for centuries. A prominent example of this regenerative power is the ability to re-grow entire amputated limbs. One important step in this regeneration process is skeletal muscle cellularization, in which the muscle fibers break down into dedifferentiated, mononuclear cells that proliferate and form new muscle in the replacement limb. In contrast, mammalian skeletal muscle does not undergo cellularization after injury. A significant proportion of research about tissue regeneration in salamanders aims to characterize regulatory genes that may have mammalian homologs. A less mainstream approach is to develop small molecule compounds that induce regeneration-related mechanisms in mammals. In this commentary, we discuss progress in discovering small molecules that induce cellularization in mammalian muscle. New research findings using these compounds has also shed light on cellular processes that regulate cellularization, such as apoptotic signaling. Although formidable technical hurdles remain, this progress increases our understanding of tissue regeneration and provide opportunities for developing small molecules that may enhance tissue repair in humans.
Harshbarger, J.C.; Chang, S.C.; DeLanney, L.E.; Rose, F.L.; Green, D.E.
Spontaneous mastocytomas studied in 18 axolotls (Ambystoma mexicanum) and six tiger salamanders (Ambystoma tigrinum) were gray-white, uni- to multilobular cutaneous protrusions from 2mm to 2cm in diameter. Tumors were moderately cellular unencapsulated masses that usually infiltrated the dermis and hypodermis with the destruction of intervening tissues. Some tumors were invading superficial bundles of the underlying skeletal muscle. Tumors consisted of mitotically active cells derived from a single lineage but showing a range of differentiation. Immature cells had nearly smooth to lightly cleft or folded basophilic nuclei bordered by a band of cytoplasm with few cytoplasmic processes and containing a few small uniform eccentric granules. Mature cells had basophilic nuclei with deep clefts or folds and abundant eosinophilic cytoplasm with multiple long intertwining cytoplasmic extensions packed with metachromatic granules. The axolotls were old individuals from an inbred laboratory colony. The tiger salamanders were wild animals from a single polluted pond. They could have been old and inbred. Both groups were neotenic. These are the first mastocytomas discovered in cold-blooded animals.
Harshbarger, J C; Chang, S C; DeLanney, L E; Rose, F L; Green, D E
Spontaneous mastocytomas studied in 18 axolotls (Ambystoma mexicanum) and six tiger salamanders (Ambystoma tigrinum) were gray-white, uni- to multilobular cutaneous protrusions from 2 mm to 2 cm in diameter. Tumors were moderately cellular unencapsulated masses that usually infiltrated the dermis and hypodermis with the destruction of intervening tissues. Some tumors were invading superficial bundles of the underlying skeletal muscle. Tumors consisted of mitotically active cells derived from a single lineage but showing a range of differentiation. Immature cells had nearly smooth to lightly cleft or folded basophilic nuclei bordered by a band of cytoplasm with few cytoplasmic processes and containing a few small uniform eccentric granules. Mature cells had basophilic nuclei with deep clefts or folds and abundant eosinophilic cytoplasm with multiple long intertwining cytoplasmic extensions packed with metachromatic granules. The axolotls were old individuals from an inbred laboratory colony. The tiger salamanders were wild animals from a single polluted pond. They could have been old and inbred. Both groups were neotenic. These are the first mastocytomas discovered in cold-blooded animals.
Woodcock, M Ryan; Vaughn-Wolfe, Jennifer; Elias, Alexandra; Kump, D Kevin; Kendall, Katharina Denise; Timoshevskaya, Nataliya; Timoshevskiy, Vladimir; Perry, Dustin W; Smith, Jeramiah J; Spiewak, Jessica E; Parichy, David M; Voss, S Randal
The molecular genetic toolkit of the Mexican axolotl, a classic model organism, has matured to the point where it is now possible to identify genes for mutant phenotypes. We used a positional cloning-candidate gene approach to identify molecular bases for two historic axolotl pigment phenotypes: white and albino. White (d/d) mutants have defects in pigment cell morphogenesis and differentiation, whereas albino (a/a) mutants lack melanin. We identified in white mutants a transcriptional defect in endothelin 3 (edn3), encoding a peptide factor that promotes pigment cell migration and differentiation in other vertebrates. Transgenic restoration of Edn3 expression rescued the homozygous white mutant phenotype. We mapped the albino locus to tyrosinase (tyr) and identified polymorphisms shared between the albino allele (tyr (a) ) and tyr alleles in a Minnesota population of tiger salamanders from which the albino trait was introgressed. tyr (a) has a 142 bp deletion and similar engineered alleles recapitulated the albino phenotype. Finally, we show that historical introgression of tyr (a) significantly altered genomic composition of the laboratory axolotl, yielding a distinct, hybrid strain of ambystomatid salamander. Our results demonstrate the feasibility of identifying genes for traits in the laboratory Mexican axolotl.
Zang, Jingjing; Matthews, Hugh R
Recovery of the light response in vertebrate photoreceptors requires the shutoff of both active intermediates in the phototransduction cascade: the visual pigment and the transducin-phosphodiesterase complex. Whichever intermediate quenches more slowly will dominate photoresponse recovery. In suction pipette recordings from isolated salamander ultraviolet- and blue-sensitive cones, response recovery was delayed, and the dominant time constant slowed when internal [Ca(2+)] was prevented from changing after a bright flash by exposure to 0Ca(2+)/0Na(+) solution. Taken together with a similar prior observation in salamander red-sensitive cones, these observations indicate that the dominance of response recovery by a Ca(2+)-sensitive process is a general feature of amphibian cone phototransduction. Moreover, changes in the external pH also influenced the dominant time constant of red-sensitive cones even when changes in internal [Ca(2+)] were prevented. Because the cone photopigment is, uniquely, exposed to the external solution, this may represent a direct effect of protons on the equilibrium between its inactive Meta I and active Meta II forms, consistent with the notion that the process dominating recovery of the bright flash response represents quenching of the active Meta II form of the cone photopigment.
Stinson, Charlotte M; Deban, Stephen M
During aquatic feeding salamanders use the hyobranchial apparatus to capture prey. The hyobranchial apparatus depresses the floor of the mouth, effectively expanding the oropharyngeal cavity and generating suction. Within the family Salamandridae, there is a wide range of ecological diversity, with salamanders being terrestrial, semi-aquatic, or aquatic as adults. The purpose of this research was to quantify the diverse morphology and suction feeding performance of aquatically feeding salamandrids. We hypothesized that a more robust hyobranchial apparatus morphology would yield increased aquatic feeding performance. When compared to semi-aquatic newts, the fully aquatic species Paramesotriton labiatus had greater mineralization of the hyobranchial apparatus, as well as relatively more narrow basibranchial and wider ceratobranchial I+II complexes. These morphological differences coincide with greater aquatic feeding performance. Kinematics from high-speed videography revealed that maximum mouth opening velocity and acceleration were approximately two and five times greater, respectively, in Paramesotriton, and hyobranchial depression acceleration was found to be approximately three times greater than in the semi-aquatic species Pleurodeles waltl, Notophthalmus viridescens, Triturus dobrogicus, and Cynops cyanurus. Using digital particle image velocimetry, peak and average fluid velocity generated in Paramesotriton during suction feeding events were found to be 0.5ms(-1) and 0.2ms(-1), respectively, doubling that of all semi-aquatic species. These findings reveal that specialized morphology increases aquatic feeding performance in a fully aquatic newt. Copyright © 2017 Elsevier GmbH. All rights reserved.
Jiang, Wen-Bin; Hakim, Ma; Luo, Lei; Li, Bo-Wen; Yang, Shi-Long; Song, Yu-Zhu; Lai, Ren; Lu, Qiu-Min
As a group of intestinal hormones and neurotransmitters, cholecystokinins (CCKs) regulate and affect pancreatic enzyme secretion, gastrointestinal motility, pain hypersensitivity, digestion and satiety, and generally contain a DYMGWMDFG sequence at the C-terminus. Many CCKs have been reported in mammals. However, only a few have been reported in amphibians, such as Hyla nigrovittata, Xenopus laevis, and Rana catesbeiana, with none reported in urodele amphibians like newts and salamanders. Here, a CCK called CCK-TV was identified and characterized from the skin of the salamander Tylototriton verrucosus. This CCK contained an amino acid sequence of DYMGWMDF-NH2 as seen in other CCKs. A cDNA encoding the CCK precursor containing 129 amino acid residues was cloned from the cDNA library of T. verrucosus skin. The CCK-TV had the potential to induce the contraction of smooth muscle strips isolated from porcine gallbladder, eliciting contraction at a concentration of 5.0 x 10⁻¹¹ mol/L and inducing maximal contraction at a concentration of 2.0 x 10⁻⁶ mol/L. The EC50 was 13.6 nmol/L. To the best of our knowledge, this is the first report to identify the presence of a CCK in an urodele amphibian.
Matsunami, Masatoshi; Kitano, Jun; Kishida, Osamu; Michimae, Hirofumi; Miura, Toru; Nishimura, Kinya
Predator- and prey-induced phenotypic plasticity is widely observed among amphibian species. Although ecological factors inducing diverse phenotypic responses have been extensively characterized, we know little about the molecular bases of variation in phenotypic plasticity. Larvae of the Hokkaido salamander, Hynobius retardatus, exhibit two distinct morphs: the presence of their prey, Rana pirica tadpoles, induces a broad-headed attack morph, and the presence of predatory dragonfly nymphs (Aeshna nigroflava) induces a defence morph with enlarged external gills and a high tail. To compare the genes involved in predator- and prey-induced phenotypic plasticity, we carried out a de novo transcriptome analysis of Hokkaido salamander larvae exposed to either prey or predator individuals. First, we found that the number of genes involved in the expression of the defence morph was approximately five times the number involved in the expression of the attack morph. This result is consistent with the fact that the predator-induced plasticity involves more drastic morphological changes than the prey-induced plasticity. Second, we found that particular sets of genes were upregulated during the induction of both the attack and defence morphs, but others were specific to the expression of one or the other morph. Because both shared and unique molecular mechanisms were used in the expression of each morph, the evolution of a new plastic phenotype might involve both the co-option of pre-existing molecular mechanisms and the acquisition of novel regulatory mechanisms.
Shen, Xing-Xing; Liang, Dan; Chen, Meng-Yun; Mao, Rong-Li; Wake, David B; Zhang, Peng
Deep phylogenetic relationships of the largest salamander family Plethodontidae have been difficult to resolve, probably reflecting a rapid diversification early in their evolutionary history. Here, data from 50 independent nuclear markers (total 48,582 bp) are used to reconstruct the phylogeny and divergence times for plethodontid salamanders, using both concatenation and coalescence-based species tree analyses. Our results robustly resolve the position of the enigmatic eastern North American four-toed salamander (Hemidactylium) as the sister taxon of Batrachoseps + Tribe Bolitoglossini, thus settling a long-standing question. Furthermore, we statistically reject sister taxon status of Karsenia and Hydromantes, the only plethodontids to occur outside the Americas, leading us to new biogeographic hypotheses. Contrary to previous long-standing arguments that plethodontid salamanders are an old lineage originating in the Cretaceous (more than 90 Ma), our analyses lead to the hypothesis that these salamanders are much younger, arising close to the K-T boundary (~66 Ma). These time estimates are highly stable using alternative calibration schemes and dating methods. Our data simulation highlights the potential risk of making strong arguments about phylogenetic timing based on inferences from a handful of nuclear genes, a common practice. Based on the newly obtained timetree and ancestral area reconstruction results, we argue that (i) the classic "Out of Appalachia" hypothesis of plethodontid origins is problematic; (ii) the common ancestor of extant plethodontids may have originated in northwestern North America in the early Paleocene; (iii) origins of Eurasian plethodontids likely result from two separate dispersal events from western North America via Beringia in the late Eocene (~42 Ma) and the early Miocene (~23 Ma), respectively.
U.S. Geological Survey, Department of the Interior — Data exist in a Microsoft Access 2003 database, and are the result of fixed-area, time-constrained searches for terrestrial salamanders within and nearby a...
Full Text Available Most amphibians use both wetland and upland habitats, but the extent of their movement in forested habitats is poorly known. We used radiotelemetry to observe the movements of adult and juvenile eastern tiger salamanders over a 4-year period. Females tended to move farther from the breeding ponds into upland forested habitat than males, while the distance a juvenile moved appeared to be related to body size, with the largest individuals moving as far as the adult females. Individuals chose refugia in native pitch pine—oak forested habitat and avoided open fields, roads, and developed areas. We also observed a difference in potential predation pressures in relation to the distance an individual moved from the edge of the pond. Our results support delineating forested wetland buffer zones on a case-by-case basis to reduce the impacts of concentrated predation, to increase and protect the availability of pitch pine—oak forests near the breeding pond, and to focus primarily on the habitat needs of the adult females and larger juveniles, which in turn will encompass habitat needs of adult males and smaller juveniles.
Moore, Jean-David; Ouellet, Martin
The effects of recent climate changes on earth ecosystems are likely among the most important ecological concerns in human history. Good bioindicators are essential to properly assess the magnitude of these changes. In the last decades, studies have suggested that the morph proportion of the eastern red-backed salamander (Plethodon cinereus), one of the most widely distributed and abundant vertebrate species in forests of eastern North America, could be used as a proxy for monitoring climate changes. Based on new discoveries in the northern areas of the species' range and on one of the largest compilation ever made for a vertebrate in North America (236 109 observations compiled from 1880 to 2013 in 1148 localities), we demonstrate however that climatic and geographic variables do not influence the colour morph proportions in P. cinereus populations. Consequently, we show that the use of colour morph proportions of this species do not perform as an indicator of climate change. Our findings indicate that bioindicator paradigms can be significantly challenged by new ecological research and more representative databases. © 2014 Her Majesty the Queen in Right of Canada Global Change Biology © 2014 John Wiley & Sons Ltd.
Thomas, Jessica R; Woodley, Sarah K
In vertebrates, exposure to stressors and stress hormones has a number of physiological effects including modulation of immune function. These effects on immune function have been well studied in mammals, but less is known in other groups, in particular amphibians. To analyze the effects of exposure to stressors and the stress hormone corticosterone, we monitored cutaneous wound healing as a measure of integrated immunity in male and female semi-terrestrial salamanders (Desmognathus ochrophaeus) that were chased to induce endogenous release of corticosterone or were treated with physiologically relevant doses of corticosterone. As predicted, subjects treated daily with corticosterone healed more slowly than did controls. In contrast, subjects that had been chased daily healed at the same rate as controls. Surprisingly, repeated chasing did not elevate plasma corticosterone despite causing drops in body mass and survival. Additionally, females healed more slowly than males, possibly due to energetic constraints.
Weizhao YANG; Jie WANG; Jianping JIANG; Cheng LI; Feng XIE
Echinotriton chinhaiensis is a critically endangered salamander and its distribution is restricted to Ningbo, Zhejiang Province of China. In this study, we developed and characterized iffteen polymorphic microsatellite loci forE. chinhaiensis from dinucleotide- and tetranucleotide- enriched library. The number of alleles ranges from 4 to 12 with an average of 7.27 alleles per locus. The observed and expected heterozygosities values were from 0.250 to 0.844 and 0.511 to 0.872 with an average value of 0.596 and 0.722; respectively. The polymorphic microsatellite loci described in this paper are useful in the further study on genetic diversity and gene lfow, which would be helpful to formulate effective conservation strategies for theE. chinhaiensis.
Ehmcke, Jens; Clemen, Günter
The skulls of six species of plethodontid salamanders of the genus Bolitoglossa from Costa Rica and Panama, Bolitoglossa colonnea, B. dofleini, B. lignicolor, B. marmorea, B. schizodactyla and B. striatula are comparatively analysed. All species are terrestrial or slightly arboreal and show no life-mode-dependent skull characteristics. Heads cleared by transparent preparation and stained in toto were used for examination of the skull structure and paraffin sections of the gonads were prepared to confirm sexual maturity of each individual. Focussing on the size of the premaxillary pars dentalis, the degree of fusion of the processus dorsales praemaxillares and the presence or absence of the paired prefrontals we state that B. dofleini and B. striatula possess more ancestral characteristics, B. marmorea is situated in an intermediate position and B. colonnea, B. lignicolor and B. schizodactyla show more advanced characteristics.
Diaz Quiroz, Juan Felipe; Echeverri, Karen
Major trauma to the mammalian spinal cord often results in irreversible loss of function, i.e. paralysis, and current therapies ranging from drugs, implantations of stem cells and/or biomaterials, and electrically stimulated nerve regrowth, have so far offered very limited success in improving quality-of-life. However, in marked contrast with this basic shortcoming of ours, certain vertebrate species, including fish and salamanders, display the amazing ability to faithfully regenerate various complex body structures after injury or ablation, restoring full functionality, even in the case of the spinal cord. Despite the inherently strong and obvious translational potential for improving treatment strategies for human patients, our in-depth molecular-level understanding of these decidedly more advanced repair systems remains in its infancy. In the present review, we will discuss the current state of this field, focusing on recent progress in such molecular analyses using various regenerative species, and how these so far relate to the mammalian situation.
Berg, Daniel A; Kirkham, Matthew; Wang, Heng; Frisén, Jonas; Simon, András
Appropriate termination of regenerative processes is critical for producing the correct number of cells in tissues. Here we provide evidence for an end-product inhibition of dopamine neuron regeneration that is mediated by dopamine. Ablation of midbrain dopamine neurons leads to complete regeneration in salamanders. Regeneration involves extensive neurogenesis and requires activation of quiescent ependymoglia cells, which express dopamine receptors. Pharmacological compensation for dopamine loss by L-dopa inhibits ependymoglia proliferation and regeneration in a dopamine receptor-signaling-dependent manner, specifically after ablation of dopamine neurons. Systemic administration of the dopamine receptor antagonist haloperidol alone causes ependymoglia proliferation and the appearance of excessive number of neurons. Our data show that stem cell quiescence is under dopamine control and provide a model for termination once normal homeostasis is restored. The findings establish a role for dopamine in the reversible suppression of neurogenesis in the midbrain and have implications for regenerative strategies in Parkinson's disease.
Macey, J. Robert
A new parsimony analysis of 27 complete mitochondrial genomic sequences is conducted to investigate the phylogenetic relationships of plethodontid salamanders. This analysis focuses on the amount of character conflict between phylogenetic trees recovered from newly conducted parsimony searches and the Bayesian and maximum likelihood topology reported by Mueller et al. (2004, PNAS, 101, 13820-13825). Strong support for Hemidactylium as the sister taxon to all other plethodontids is recovered from parsimony analyses. Plotting area relationships on the most parsimonious phylogenetic tree suggests that eastern North America is the origin of the family Plethodontidae supporting the ''Out of Appalachia'' hypothesis. A new taxonomy that recognizes clades recovered from phylogenetic analyses is proposed.
The Ezo salamander, Hynobius retardatus, is endemic only to Hokkaido, the northernmost island of Japan. Gravid flukes of the family Gorgoderidae were discovered from the urinary bladder of H. retardatus. The parasites were identified as a new species named Phyllodistomum kanae sp. nov. In the neighboring Honshu island another bladder fluke, Phyllodistomum patellare, has already been found from the Japanese newt. The new species clearly differs from P. patellare in having a spherical ovary and very weakly lobed testes. The discovery of species of Phyllodistomum from urodelan amphibians is very uncommon in Eurasia. A molecular phylogeny based on 28S ribosomal DNA suggests that sphaeriid bivalves may serve as the first intermediate host for the new species. Copyright © 2015. Published by Elsevier Ireland Ltd.
Rieke, F; Schwartz, E A
1. We have studied exocytosis and endocytosis in the synaptic terminal of salamander rods using a combination of Ca2+ imaging, capacitance measurement and the photolysis of Ca2+ buffers. 2. The average cytoplasmic Ca2+ concentration at the dark resting potential was 2-4 microM. 3. An average cytoplasmic Ca2+ concentration of 2-4 microM maintained a high rate of continuous exocytosis and endocytosis. 4. Changes in the rate of exocytosis were followed in less than 0.7 s by compensatory changes in the rate of endocytosis. 5. Vesicle cycling in the rod synapse is specialized for graded transmission and differs from that previously described for synapses that release synchronized bursts of transmitter.
Ficetola, Gentile Francesco; Colleoni, Emiliano; Renaud, Julien; Scali, Stefano; Padoa-Schioppa, Emilio; Thuiller, Wilfried
Despite the recognition that some species might quickly adapt to new conditions under climate change, demonstrating and predicting such a fundamental response is challenging. Morphological variations in response to climate may be caused by evolutionary changes or phenotypic plasticity, or both, but teasing apart these processes is difficult. Here, we built on the number of thoracic vertebrae (NTV) in ectothermic vertebrates, a known genetically based feature, to establish a link with body size and evaluate how climate change might affect the future morphological response of this group of species. First, we show that in old-world salamanders, NTV variation is strongly related to changes in body size. Secondly, using 22 salamander species as a case study, we found support for relationships between the spatial variation in selected bioclimatic variables and NTV for most of species. For 44% of species, precipitation and aridity were the predominant drivers of geographical variation of the NTV. Temperature features were dominant for 31% of species, while for 19% temperature and precipitation played a comparable role. This two-step analysis demonstrates that ectothermic vertebrates may evolve in response to climate change by modifying the number of thoracic vertebrae. These findings allow to develop scenarios for potential morphological evolution under future climate change and to identify areas and species in which the most marked evolutionary responses are expected. Resistance to climate change estimated from species distribution models was positively related to present-day species morphological response, suggesting that the ability of morphological evolution may play a role for species' persistence under climate change. The possibility that present-day capacity for local adaptation might help the resistance response to climate change can be integrated into analyses of the impact of global changes and should also be considered when planning management actions favouring
Ultsch, Gordon R
The giant salamanders are aquatic and paedomorphic urodeles including the genera Andrias and Cryptobranchus (Cryptobranchidae), Amphiuma (Amphiumidae), Siren (Sirenidae), and Necturus (Proteidae, of which only N. maculosus is considered 'a giant'). Species in the genera Cryptobranchus and Necturus are considered aquatic salamanders well adapted for breathing water, poorly adapted for breathing air, and with limited abilities to compensate acid-base disturbances. As such, they are water-breathing animals with a somewhat fish-like respiratory and acid-base physiology, whose habitat selection is limited to waters that do not typically become hypoxic or hypercarbic (although this assertion has been questioned for N. maculosus). Siren and Amphiuma species, by contrast, are dependent upon air-breathing, have excellent lungs, inefficient (Siren) or no (Amphiuma) gills, and are obligate air-breathers with an acid-base status more similar to that of terrestrial tetrapods. As such, they can be considered to be air-breathing animals that live in water. Their response to the aquatic hypercarbia that they often encounter is to maintain intracellular pH (pH(i) ) and abandon extracellular pH regulation, a process that has been referred to as preferential pH(i) regulation. The acid-base status of some present-day tropical air-breathing fishes, and of Siren and Amphiuma, suggests that the acid-base transition from a low PCO(2) -low  system typical of water-breathing fishes to the high PCO(2) -high  systems of terrestrial tetrapods may have been completed before emergence onto land, and likely occurred in habitats that were typically both hypoxic and hypercarbic.
Reilly, Sean B; Marks, Sharyn B; Jennings, W Bryan
The accurate delimitation of evolutionary population units represents an important component in phylogeographic and conservation genetic studies. Here, we used a combined population assignment and historical demographic approach to study a complex of ecomorphologically distinctive populations of Black Salamanders (Aneides flavipunctatus) that are parapatrically distributed and meet at a three-way contact zone in north-western California. We used mitochondrial tree-based and multilocus clustering methods to evaluate a priori two- (Northern and Southern) and three (Northern, Coast and Inland) population hypotheses derived from previous studies. Mitochondrial results were consistent with the two- and three-population hypotheses, while the nDNA clustering results supported only the two-population hypothesis. Historical demographic analyses and mtDNA gene divergence estimates revealed that the Northern and Southern populations split during the Pliocene (2-5 Ma). Subdivision of the Southern population into Coast and Inland populations was estimated to be late Pleistocene (0.24 Ma), although our mtDNA results suggested a Pliocene divergence. Effective gene flow estimates (2N(e)m) suggest that either the two- or three-population hypotheses remain valid. However, our results unexpectedly revealed that the Northern population might instead represent two parapatric populations that separated nearly 4 Ma. These results are surprising because the Pliocene divergence between these ecomorphologically conservative forms is similar or older than for the ecomorphologically divergent Coast and Inland sister populations. We conclude that Black Salamanders in north-western California belong to at least three or four populations or species, and these all meet criteria for being Evolutionary Significant Units or 'ESUs' and therefore warrant conservation consideration.
Scott, David E; Komoroski, Mark J; Croshaw, Dean A; Dixon, Philip M
Terrestrial habitats surrounding isolated wetlands are a critical resource for many pond-breeding amphibian species, yet few studies have examined the terrestrial distribution of post-metamorphic juveniles and adults. We used an encircling drift fence at a breeding pond in conjunction with partial fences at 90, 172, and 332 m from the wetland to estimate the terrestrial distribution of adult marbled salamanders (Ambystoma opacum; four breeding seasons) and mole salamanders (A. talpoideum; two seasons), as well as the dispersion of newly metamorphosed A. opacum (one summer). For newly metamorphosed A. opacum, 79% emigrated < 90 m from the wetland, and 8% moved beyond 172 m; movement distance was unrelated to body size. Distribution of adult A. opacum varied among years, with an average of 28% (range 23-31%) occurring beyond 172 m in all years. Averaged across two years, 51% of adult A. talpoideum occurred beyond 172 m. Lognormal models provided a good fit to both the juvenile and adult ambystomatid distributions, and parameters differed between age classes, sexes, species, and years within species. For adult A. opacum a buffer radius of 300 m or 340 m, depending on the year, is estimated to include 95% of adults; for A. talpoideum the estimate is 464 m or 501 m. A reanalysis of distribution data for seven ambystomatid species shows that a previous estimate of a 164-m radius to protect 95% of a population underestimates the needed buffer radius by 185 m. Because our study wetland requires a nearly 500 m wide radius to protect 95% of its ambystomatid adults, preservation of similar communities may require much more surrounding terrestrial habitat than previously thought.
Full Text Available Abstract Background Although some mechanisms of habitat adaptation of conspecific populations have been recently elucidated, the evolution of female preference has rarely been addressed as a force driving habitat adaptation in natural settings. Habitat adaptation of fire salamanders (Salamandra salamandra, as found in Middle Europe (Germany, can be framed in an explicit phylogeographic framework that allows for the evolution of habitat adaptation between distinct populations to be traced. Typically, females of S. salamandra only deposit their larvae in small permanent streams. However, some populations of the western post-glacial recolonization lineage use small temporary ponds as larval habitats. Pond larvae display several habitat-specific adaptations that are absent in stream-adapted larvae. We conducted mate preference tests with females from three distinct German populations in order to determine the influence of habitat adaptation versus neutral genetic distance on female mate choice. Two populations that we tested belong to the western post-glacial recolonization group, but are adapted to either stream or pond habitats. The third population is adapted to streams but represents the eastern recolonization lineage. Results Despite large genetic distances with FST values around 0.5, the stream-adapted females preferred males from the same habitat type regardless of genetic distance. Conversely, pond-adapted females did not prefer males from their own population when compared to stream-adapted individuals of either lineage. Conclusion A comparative analysis of our data showed that habitat adaptation rather than neutral genetic distance correlates with female preference in these salamanders, and that habitat-dependent female preference of a specific pond-reproducing population may have been lost during adaptation to the novel environmental conditions of ponds.
Bryant Susan V
Full Text Available Abstract Background Establishing genomic resources for closely related species will provide comparative insights that are crucial for understanding diversity and variability at multiple levels of biological organization. We developed ESTs for Mexican axolotl (Ambystoma mexicanum and Eastern tiger salamander (A. tigrinum tigrinum, species with deep and diverse research histories. Results Approximately 40,000 quality cDNA sequences were isolated for these species from various tissues, including regenerating limb and tail. These sequences and an existing set of 16,030 cDNA sequences for A. mexicanum were processed to yield 35,413 and 20,599 high quality ESTs for A. mexicanum and A. t. tigrinum, respectively. Because the A. t. tigrinum ESTs were obtained primarily from a normalized library, an approximately equal number of contigs were obtained for each species, with 21,091 unique contigs identified overall. The 10,592 contigs that showed significant similarity to sequences from the human RefSeq database reflected a diverse array of molecular functions and biological processes, with many corresponding to genes expressed during spinal cord injury in rat and fin regeneration in zebrafish. To demonstrate the utility of these EST resources, we searched databases to identify probes for regeneration research, characterized intra- and interspecific nucleotide polymorphism, saturated a human – Ambystoma synteny group with marker loci, and extended PCR primer sets designed for A. mexicanum / A. t. tigrinum orthologues to a related tiger salamander species. Conclusions Our study highlights the value of developing resources in traditional model systems where the likelihood of information transfer to multiple, closely related taxa is high, thus simultaneously enabling both laboratory and natural history research.
Chong, Rebecca A; Mueller, Rachel Lockridge
Mitochondria are intracellular organelles where oxidative phosphorylation is carried out to complete ATP synthesis. Mitochondria have their own genome; in metazoans, this is a small, circular molecule encoding 13 electron transport proteins, 22 tRNAs, and 2 rRNAs. In invertebrates, mitochondrial gene rearrangement is common, and it is correlated with increased substitution rates. In vertebrates, mitochondrial gene rearrangement is rare, and its relationship to substitution rate remains unexplored. Mitochondrial genes can also show spatial variation in substitution rates around the genome due to the mechanism of mtDNA replication, which produces a mutation gradient. To date, however, the strength of the mutation gradient and whether movement along the gradient in rearranged (or otherwise modified) genomes impacts genic substitution rates remain unexplored in the majority of vertebrates. Salamanders include both normal mitochondrial genomes and independently derived rearrangements and expansions, providing a rare opportunity to test the effects of large-scale changes to genome architecture on vertebrate mitochondrial gene sequence evolution. We show that: 1) rearranged/expanded genomes have higher substitution rates; 2) most genes in rearranged/expanded genomes maintain their position along the mutation gradient, substitution rates of the genes that do move are unaffected by their new position, and the gradient in salamanders is weak; and 3) genomic rearrangements/expansions occur independent of levels of selective constraint on genes. Together, our results demonstrate that large-scale changes to genome architecture impact mitochondrial gene evolution in predictable ways; however, despite these impacts, the same functional constraints act on mitochondrial protein-coding genes in both modified and normal genomes.
Kono, Masahiro; Crouch, Rosalie K; Oprian, Daniel D
A triple mutant (F86L/T93P/S118T; bovine rhodopsin numbering) of the tiger salamander UV cone pigment appears to be trapped in an open conformation that is metarhodopsin-II-like. The pigment is able to activate transducin in the dark, and the ligand-free apoprotein is also able to activate transducin constitutively. The pigment permits protons and chloride ions from solution access to the active site as it displays a pH- and NaCl-dependent absorption spectrum not observed with the wild-type pigment. However, the wild-type properties of light-dependent activity and a pH-independent absorption spectrum are recovered upon reconstitution of the triple mutant with 11-cis-9-demethyl retinal. These results suggest that binding the native chromophore cannot deactivate the protein because of steric interactions between the protein, possibly residue 118, and the 9-methyl group of the chromophore. Furthermore, the absorption spectrum of the 9-demethyl retinal regenerated pigment exhibits a band broader and with lower extinction at the absorption maximum than either the human blue or salamander UV wild-type pigments generated with the same retinal analogue. The broad spectrum appears to be comprised of two or more species and can be well-fit by a sum of scaled spectra of the two wild-type pigments. Binding the chromophore appears to trap the pigment in two or more conformations. The triple mutant reported here represents the first example of a dark-active cone pigment and constitutively active cone opsin.
Gareth J Voss
Full Text Available Very little is known about the factors that cause variation in regenerative potential within and between species. Here, we used a genetic approach to identify heritable genetic factors that explain variation in tail regenerative outgrowth. A hybrid ambystomatid salamander (Ambystoma mexicanum x A. andersoni was crossed to an A. mexicanum and 217 offspring were induced to undergo metamorphosis and attain terrestrial adult morphology using thyroid hormone. Following metamorphosis, each salamander's tail tip was amputated and allowed to regenerate, and then amputated a second time and allowed to regenerate. Also, DNA was isolated from all individuals and genotypes were determined for 187 molecular markers distributed throughout the genome. The area of tissue that regenerated after the first and second amputations was highly positively correlated across males and females. Males presented wider tails and regenerated more tail tissue during both episodes of regeneration. Approximately 66-68% of the variation in regenerative outgrowth was explained by tail width, while tail length and genetic sex did not explain a significant amount of variation. A small effect QTL was identified as having a sex-independent effect on tail regeneration, but this QTL was only identified for the first episode of regeneration. Several molecular markers significantly affected regenerative outgrowth during both episodes of regeneration, but the effect sizes were small (<4% and correlated with tail width. The results show that ambysex and minor effect QTL explain variation in adult tail morphology and importantly, tail width. In turn, tail width at the amputation plane largely determines the rate of regenerative outgrowth. Because amputations in this study were made at approximately the same position of the tail, our results resolve an outstanding question in regenerative biology: regenerative outgrowth positively co-varies as a function of tail width at the amputation site.
Voss, Gareth J; Kump, D Kevin; Walker, John A; Voss, S Randal
Very little is known about the factors that cause variation in regenerative potential within and between species. Here, we used a genetic approach to identify heritable genetic factors that explain variation in tail regenerative outgrowth. A hybrid ambystomatid salamander (Ambystoma mexicanum x A. andersoni) was crossed to an A. mexicanum and 217 offspring were induced to undergo metamorphosis and attain terrestrial adult morphology using thyroid hormone. Following metamorphosis, each salamander's tail tip was amputated and allowed to regenerate, and then amputated a second time and allowed to regenerate. Also, DNA was isolated from all individuals and genotypes were determined for 187 molecular markers distributed throughout the genome. The area of tissue that regenerated after the first and second amputations was highly positively correlated across males and females. Males presented wider tails and regenerated more tail tissue during both episodes of regeneration. Approximately 66-68% of the variation in regenerative outgrowth was explained by tail width, while tail length and genetic sex did not explain a significant amount of variation. A small effect QTL was identified as having a sex-independent effect on tail regeneration, but this QTL was only identified for the first episode of regeneration. Several molecular markers significantly affected regenerative outgrowth during both episodes of regeneration, but the effect sizes were small (<4%) and correlated with tail width. The results show that ambysex and minor effect QTL explain variation in adult tail morphology and importantly, tail width. In turn, tail width at the amputation plane largely determines the rate of regenerative outgrowth. Because amputations in this study were made at approximately the same position of the tail, our results resolve an outstanding question in regenerative biology: regenerative outgrowth positively co-varies as a function of tail width at the amputation site.
Vaglia, Janet L; Fornari, Chet; Evans, Paula K
During embryogenesis, the body axis elongates and specializes. In vertebrate groups such as salamanders and lizards, elongation of the posterior body axis (tail) continues throughout life. This phenomenon of post-embryonic tail elongation via addition of vertebrae has remained largely unexplored, and little is known about the underlying developmental mechanisms that promote vertebral addition. Our research investigated tail elongation across life stages in a non-model salamander species, Eurycea cirrigera (Plethodontidae). Post-embryonic addition of segments suggests that the tail tip retains some aspects of embryonic cell/tissue organization and gene expression throughout the life cycle. We describe cell and tissue differentiation and segmentation of the posterior tail using serial histology and expression of the axial tissue markers, MF-20 and Pax6. Embryonic expression patterns of HoxA13 and C13 are shown with in situ hybridization. Tissue sections reveal that the posterior spinal cord forms via cavitation and precedes development of the underlying cartilaginous rod after embryogenesis. Post-embryonic tail elongation occurs in the absence of somites and mesenchymal cells lateral to the midline express MF-20. Pax6 expression was observed only in the spinal cord and some mesenchymal cells of adult Eurycea tails. Distinct temporal and spatial patterns of posterior Hox13 gene expression were observed throughout embryogenesis. Overall, important insights to cell organization, differentiation, and posterior Hox gene expression may be gained from this work. We suggest that further work on gene expression in the elongating adult tail could shed light on mechanisms that link continual axial elongation with regeneration.
Putta, Srikrishna; Smith, Jeramiah J; Walker, John A; Rondet, Mathieu; Weisrock, David W; Monaghan, James; Samuels, Amy K; Kump, Kevin; King, David C; Maness, Nicholas J; Habermann, Bianca; Tanaka, Elly; Bryant, Susan V; Gardiner, David M; Parichy, David M; Voss, S Randal
Background Establishing genomic resources for closely related species will provide comparative insights that are crucial for understanding diversity and variability at multiple levels of biological organization. We developed ESTs for Mexican axolotl (Ambystoma mexicanum) and Eastern tiger salamander (A. tigrinum tigrinum), species with deep and diverse research histories. Results Approximately 40,000 quality cDNA sequences were isolated for these species from various tissues, including regenerating limb and tail. These sequences and an existing set of 16,030 cDNA sequences for A. mexicanum were processed to yield 35,413 and 20,599 high quality ESTs for A. mexicanum and A. t. tigrinum, respectively. Because the A. t. tigrinum ESTs were obtained primarily from a normalized library, an approximately equal number of contigs were obtained for each species, with 21,091 unique contigs identified overall. The 10,592 contigs that showed significant similarity to sequences from the human RefSeq database reflected a diverse array of molecular functions and biological processes, with many corresponding to genes expressed during spinal cord injury in rat and fin regeneration in zebrafish. To demonstrate the utility of these EST resources, we searched databases to identify probes for regeneration research, characterized intra- and interspecific nucleotide polymorphism, saturated a human – Ambystoma synteny group with marker loci, and extended PCR primer sets designed for A. mexicanum / A. t. tigrinum orthologues to a related tiger salamander species. Conclusions Our study highlights the value of developing resources in traditional model systems where the likelihood of information transfer to multiple, closely related taxa is high, thus simultaneously enabling both laboratory and natural history research. PMID:15310388
Zhu, Wei; Kuo, Dwight; Nathanson, Jason; Satoh, Akira; Pao, Gerald M.; Yeo, Gene W.; Bryant, Susan V.; Voss, S. Randal; Gardiner, David M.; Hunter, Tony
Salamanders possess an extraordinary capacity for tissue and organ regeneration when compared to mammals. In our effort to characterize the unique transcriptional fingerprint emerging during the early phase of salamander limb regeneration, we identified transcriptional activation of some germline-specific genes within the Mexican axolotl (Ambystoma mexicanum) that is indicative of cellular reprogramming of differentiated cells into a germline-like state. In this work, we focus on one of these genes, the long interspersed nucleotide element-1 (LINE-1) retrotransposon, which is usually active in germ cells and silent in most of the somatic tissues in other organisms. LINE-1 was found to be dramatically upregulated during regeneration. In addition, higher genomic LINE-1 content was also detected in the limb regenerate when compared to that before amputation indicating that LINE-1 retrotransposition is indeed active during regeneration. Active LINE-1 retrotransposition has been suggested to have a potentially deleterious impact on genomic integrity. Silencing of activated LINE-1 by small RNAs has been reported to be part of the machinery aiming to maintain genomic integrity. Indeed, we were able to identify putative LINE-1-related piRNAs in the limb blastema. Transposable element-related piRNAs have been identified frequently in the germline in other organisms. Thus, we present here a scenario in which a unique germline-like state is established during axolotl limb regeneration, and the re-activation of LINE-1 may serve as a marker for cellular dedifferentiation in the early-stage of limb regeneration. PMID:22913491
Chelgren, N.D.; Adams, M.J.; Bailey, L.L.; Bury, R.B.
Studies of the distribution of elusive forest wildlife have suffered from the confounding of true presence with the uncertainty of detection. Occupancy modeling, which incorporates probabilities of species detection conditional on presence, is an emerging approach for reducing observation bias. However, the current likelihood modeling framework is restrictive for handling unexplained sources of variation in the response that may occur when there are dependence structures such as smaller sampling units that are nested within larger sampling units. We used multilevel Bayesian occupancy modeling to handle dependence structures and to partition sources of variation in occupancy of sites by terrestrial salamanders (family Plethodontidae) within and surrounding an earlier wildfire in western Oregon, USA. Comparison of model fit favored a spatial N-mixture model that accounted for variation in salamander abundance over models that were based on binary detection/non-detection data. Though catch per unit effort was higher in burned areas than unburned, there was strong support that this pattern was due to a higher probability of capture for individuals in burned plots. Within the burn, the odds of capturing an individual given it was present were 2.06 times the odds outside the burn, reflecting reduced complexity of ground cover in the burn. There was weak support that true occupancy was lower within the burned area. While the odds of occupancy in the burn were 0.49 times the odds outside the burn among the five species, the magnitude of variation attributed to the burn was small in comparison to variation attributed to other landscape variables and to unexplained, spatially autocorrelated random variation. While ordinary occupancy models may separate the biological pattern of interest from variation in detection probability when all sources of variation are known, the addition of random effects structures for unexplained sources of variation in occupancy and detection
Makino, C L; Kraft, T W; Mathies, R A; Lugtenburg, J; Miley, M E; van der Steen, R; Baylor, D A
1. Chemically modified retinal chromophores were used to investigate the mechanisms that produce the characteristic spectral absorptions of cone pigments. Spectral sensitivities of single cones from the salamander, squirrel and macaque retina were determined by electrical recording. The chromophore was then replaced by bleaching the pigment and regenerating it with a retinal analogue. 2. Exposing a bleached cone to 9-cis-retinal for a brief period (less than 20 min) caused its flash sensitivity to recover to about 0.2 of the pre-bleach value. Similar exposure to a locked 6-s-cis, 9-cis analogue gave a recovery to about 0.03 of the pre-bleach value. 3. Unlike the flash sensitivity, the saturating photocurrent amplitude often recovered completely after bleaching and regenerating the pigment. 4. When the 3-dehydroretinal chromophore in the salamander long-wavelength-sensitive (red) cone was replaced with 11-cis-retinal, shortening the conjugated chain in the chromophore, the spectral sensitivity underwent a blue shift of 67 nm. 5. Pigments containing the planar-locked 6-s-cis.9-cis-retinal analogue absorbed at substantially longer wavelength than those containing unmodified 9-cis-retinal. The opsin shift, a measure of the protein's ability to modify the chromophore's absorption was larger for the locked analogue than for 9-cis-retinal. This suggests that the native chromophore assumes a twisted 6-s-cis conformation in these pigments. 6. The spectral sensitivities of red and green macaque cones containing 9-cis-retinal or planar-locked 6-s-cis.9-cis-retinal retained the 30 nm separation characteristic of the native pigments. This suggests that the different absorptions of of the 6-7 carbon bond in the retinal chromophore.
Palmer, Catherine A; Watts, Richard A; Hastings, Amy P; Houck, Lynne D; Arnold, Stevan J
Sexual communication in plethodontid salamanders is mediated by a proteinaceous pheromone that a male delivers to a female during courtship, boosting her receptivity. The pheromone consists of three proteins from three unrelated protein families. These proteins are among a small group of pheromones known to affect female receptivity in vertebrates. Previously, we showed that the genes of two of these proteins (PRF and SPF) are prone to incessant evolution driven by positive selection, presumably as a consequence of coevolution with female receptors. In this report, we focus on the evolution of the third pheromone protein gene family, plethodontid modulating factor (PMF), to determine whether it shows the same pattern of diversification. We used RT-PCR in mental gland cDNA to survey PMF sequences from three genera of plethodontid salamanders (27 spp.) to measure rates of evolution, level of gene diversification, modes of selection, and types of amino acid substitution. Like PRF and SPF, PMF is produced by a multigene family characterized by gene duplication and high levels of polymorphism. PMF evolution is rapid, incessant, and driven by positive selection. PMF is more extreme in these dimensions than both PRF and SPF. Nestled within this extraordinary variation, however, is a signature of purifying selection, acting to preserve important structural and biochemical features of the PMF protein (i.e., secretion signal, cysteine residues, and pI). Although a pattern of persistent diversification exists at the molecular level, the morphological and behavioral aspects of the pheromone delivery system show evolutionary stasis over millions of years.
Searcy, Christopher A; Shaffer, H Bradley
Current conservation mitigation plans often fail to ensure full in-kind habitat replacement for endangered species, which suggests the need for improved methods for calculating mitigation credits. A simple, yet biologically meaningful method for calculating mitigation credits would be to let the number of mitigation credits assigned to a parcel of land scale with the reproductive value of the individuals occupying that parcel. This can be accomplished by dividing the population into 2 or more subdivisions with different reproductive values, calculating the densities of these subdivisions as a function of one or more habitat parameters, and then forming a weighted sum of these densities such that each density distribution is weighted by the reproductive value of its respective subdivision of the population. This weighted sum is the density distribution of reproductive value, and by integrating it over a particular parcel, one can determine the mitigation value of that parcel. We carried out this procedure for a population of California tiger salamanders (Ambystoma californiense), with distance from breeding site as our habitat parameter and the 3 visually identifiable age classes (adults, juveniles, and metamorphs) as our population subdivisions. This led to a density distribution of reproductive value that decreased exponentially with increasing distance from a breeding site. Mitigation strategies derived from this function will be more likely to ensure the persistence of California tiger salamander populations than current approaches, which assign all land within 1.6 km of a breeding site the same mitigation value. Use of the density distribution of reproductive value as a basis for mitigation plans is a procedure that can be applied to all endangered species, and it should improve the quality of mitigation decisions.
Ficetola, Gentile Francesco; Colleoni, Emiliano; Renaud, Julien; Scali, Stefano; Padoa-Schioppa, Emilio; Thuiller, Wilfried
Despite the recognition that some species might quickly adapt to new conditions under climate change, demonstrating and predicting such a fundamental response is challenging. Morphological variations in response to climate may be caused by evolutionary changes or phenotypic plasticity, or both, but teasing apart these processes is difficult. Here we built on the number of thoracic vertebrae (NTV) in ectothermic vertebrates, a known genetically-based feature, to establish a link with body size and evaluate how climate change might affect the future morphological response of this group of species. First we show that in old-world salamanders, NTV variation is strongly related to changes in body size. Secondly, using 22 salamander species as a case study, we found support for relationships between the spatial variation in selected bioclimatic variables and NTV for most of species. For 44% of species, precipitation and aridity were the predominant drivers of geographical variation of the NTV. Temperature features were dominant for 31% of species, while for 19% temperature and precipitation played a comparable role. This two-step analysis demonstrates that ectothermic vertebrates may evolve in response to climate change by modifying the number of thoracic vertebrae. These findings allow to develop scenarios for potential morphological evolution under future climate change, and to identify areas and species in which the most marked evolutionary responses are expected. Resistance to climate change estimated from species distribution models was positively related to present-day species morphological response, suggesting that the ability of morphological evolution may play a role for species’ persistence under climate change. The possibility that present-day capacity for local adaptation might help the resistance response to climate change can be integrated into analyses of the impact of global changes, and should also be considered when planning management actions
Jianli XIONG; Xiuying LIU; Xiaomei ZHANG; Mengyun LI; Yao MIN
Sexual dimorphism in size and shape is common in many organisms, and is a key evolutionary feature. In this study, we analyzed morphometric data of the Jilin clawed salamanderOnychodactylus zhangyapingi, an endemic Chinese salamander, to examine sexual size and shape dimorphism. The morphometric data included 14 characteristics of 13 females and 11 males and was analyzed using univariate and multivariate methods. Our results showed that sexual dimorphism occurs not only in body size, but also in body shape. Males have a longer snout-vent length than females, a rarely reported pattern of male-biased sexual size dimorphism. Females have a larger space between the axilla and groin than males, while males have longer and larger tails compared to females. The sexual dimorphism in body size and shape can be explained by existing theories, but there is little data for the mating system, behavior, reproduction, or ecology ofO. zhangyapingi, so further studies are required.
徐达宇; 雷明智; 周赟; 彭海蓉
The manufacturing technology of luggage leather with giant salamander skin was introduced. The finished leather is soft and flexible with natural grains and primitive styles. It shows that after manufacture, the giant salamander skin could be made into high value added luggage leather.%利用大鲵皮试制出的箱包革等样品，既保存了大鲵皮的天然粒面和原始风格，又具有一定的柔软度和丰满性。试验表明大鲵皮经过科学加工，能够制造出具有较高经济价值的高档箱包皮革。
Qinghua Luo; Ying Liu; Liyun Zhang
Chinese giant salamander (Andrias davidianus) is a native amphibian to China. It was listed as a Class II protected wildlife species under the state conservation law largely due to its sharply declined population size in the past two decades. Habitat destruction, habitat degradation and human over-exploitation have been considered as the primary causes of the declination. Designed to boost the wild population, a captive breeding program has been successfully established and artificially bred ...
邓捷; 王中乾; 张红星; 姜维; 王启军; 赵虎
中国大鲵是国家二级保护动物，具有很高的药用价值和特殊的进化地位，人工繁育和养殖大鲵具有科学研究和经济学双重意义。但多种高传染、高致死性疾病成为大鲵养殖业发展的瓶颈，这些疾病的病原体包括细菌、真菌、寄生虫和病毒等。本文就目前大鲵多种病害的发病原因、临床特征及相应的治疗方法进行简单综述，为中国大鲵疾病的深入研究提供必备的科学依据。%Chinese giant salamander is one of the State Second-class Protection species ,with high me‐dicinal value and evolutionary significance .Artificial breeding giant salamander has double meaning in both science research and economics .However ,a variety of highly infectious ,lethal diseases have be‐come the bottleneck in breeding industry development of giant salamander .Pathogens of these diseases are included bacteria ,fungi ,parasites and viruses .T his article briefly review s the current various dis‐eases etiology ,clinical features and appropriate treatments ,in order to provide the necessary scientific basis for in-depth study of Chinese giant salamander disease .
Maximina H. Yun; Phillip B. Gates; Jeremy P. Brockes
Summary In regeneration-competent vertebrates, such as salamanders, regeneration depends on the ability of various differentiated adult cell types to undergo natural reprogramming. This ability is rarely observed in regeneration-incompetent species such as mammals, providing an explanation for their poor regenerative potential. To date, little is known about the molecular mechanisms mediating natural reprogramming during regeneration. Here, we have identified the extent of extracellular signa...
Gandhi, Jaina S; Cecala, Kristen K
The objective of the present study was to evaluate the potential interactive effects of stream temperatures and environmentally relevant glyphosate-based herbicide concentrations on movement and antipredator behaviors of larval Eurycea wilderae (Blue Ridge two-lined salamander). Larval salamanders were exposed to 1 of 4 environmentally relevant glyphosate concentrations (0.00 µg acid equivalent [a.e.]/L, 0.73 µg a.e./L, 1.46 µg a.e./L, and 2.92 µg a.e./L) at either ambient (12 °C) or elevated (23 °C) water temperature. Behaviors observed included the exploration of a novel habitat, use of refuge, habitat selection relative to a potential predator, and burst movement distance. In the absence of glyphosate, temperature consistently affected movement and refuge-use behavior, with individuals moving longer distances more frequently and using refuge less at warm temperatures; however, when glyphosate was added, the authors observed inconsistent effects of temperature that may have resulted from differential toxicity at various temperatures. Larval salamanders made shorter, more frequent movements and demonstrated reduced burst distance at higher glyphosate concentrations. The authors also found that lower glyphosate concentrations sometimes had stronger effects than higher concentrations (i.e., nonmonotonic dose responses), suggesting that standard safety tests conducted only at higher glyphosate concentrations might overlook important sublethal effects on salamander behavior. These data demonstrate that sublethal effects of glyphosate-based herbicides on natural behaviors of amphibians can occur with short-term exposure to environmentally relevant concentrations. Environ Toxicol Chem 2016;35:2297-2303. © 2016 SETAC.
Full Text Available Here, we use a three-dimensional, neuro-musculo-mechanical model of a salamander with realistic physical parameters in order to investigate the role of sensory feedback in gait generation and transition. Activation of limb and axial muscles were driven by neural output patterns obtained from a central pattern generator (CPG which is composed of simulated spiking neurons with adaptation. The CPG consists of a body CPG and four limb CPGs that are interconnected via synapses both ipsilateraly and contralaterally. We use the model both with and without sensory modulation and for different combinations of ipsilateral and contralateral coupling between the limb CPGs. We found that the proprioceptive sensory inputs are essential in obtaining a coordinated walking gait. The sensory feedback includes the signals coming from the stretch receptor like intraspinal neurons located in the girdle regions and the limb stretch receptors residing in the hip and scapula regions of the salamander. On the other hand, coordinated motor output patterns for the trotting gait were obtainable without the sensory inputs. We found that the gait transition from walking to trotting can be induced by increased activity of the descending drive coming from the mesencephalic locomotor region (MLR and is helped by the sensory inputs at the hip and scapula regions detecting the late stance phase. More neurophysiological experiments are required to identify the precise type of mechanoreceptors in the salamander and the neural mechanisms mediating the sensory modulation.
Maximina H. Yun
Full Text Available In regeneration-competent vertebrates, such as salamanders, regeneration depends on the ability of various differentiated adult cell types to undergo natural reprogramming. This ability is rarely observed in regeneration-incompetent species such as mammals, providing an explanation for their poor regenerative potential. To date, little is known about the molecular mechanisms mediating natural reprogramming during regeneration. Here, we have identified the extent of extracellular signal-regulated kinase (ERK activation as a key component of such mechanisms. We show that sustained ERK activation following serum induction is required for re-entry into the cell cycle of postmitotic salamander muscle cells, partially by promoting the downregulation of p53 activity. Moreover, ERK activation induces epigenetic modifications and downregulation of muscle-specific genes such as Sox6. Remarkably, while long-term ERK activation is found in salamander myotubes, only transient activation is seen in their mammalian counterparts, suggesting that the extent of ERK activation could underlie differences in regenerative competence between species.
Yun, Maximina H; Gates, Phillip B; Brockes, Jeremy P
In regeneration-competent vertebrates, such as salamanders, regeneration depends on the ability of various differentiated adult cell types to undergo natural reprogramming. This ability is rarely observed in regeneration-incompetent species such as mammals, providing an explanation for their poor regenerative potential. To date, little is known about the molecular mechanisms mediating natural reprogramming during regeneration. Here, we have identified the extent of extracellular signal-regulated kinase (ERK) activation as a key component of such mechanisms. We show that sustained ERK activation following serum induction is required for re-entry into the cell cycle of postmitotic salamander muscle cells, partially by promoting the downregulation of p53 activity. Moreover, ERK activation induces epigenetic modifications and downregulation of muscle-specific genes such as Sox6. Remarkably, while long-term ERK activation is found in salamander myotubes, only transient activation is seen in their mammalian counterparts, suggesting that the extent of ERK activation could underlie differences in regenerative competence between species.
Page, Robert B; Monaghan, James R; Samuels, Amy K; Smith, Jeramiah J; Beachy, Christopher K; Voss, S Randal
Ambystomatid salamanders offer several advantages for endocrine disruption research, including genomic and bioinformatics resources, an accessible laboratory model (Ambystoma mexicanum), and natural lineages that are broadly distributed among North American habitats. We used microarray analysis to measure the relative abundance of transcripts isolated from A. mexicanum epidermis (skin) after exogenous application of thyroid hormone (TH). Only one gene had a >2-fold change in transcript abundance after 2 days of TH treatment. However, hundreds of genes showed significantly different transcript levels at days 12 and 28 in comparison to day 0. A list of 123 TH-responsive genes was identified using statistical, BLAST, and fold level criteria. Cluster analysis identified two groups of genes with similar transcription patterns: up-regulated versus down-regulated. Most notably, several keratins exhibited dramatic (1000 fold) increases or decreases in transcript abundance. Keratin gene expression changes coincided with morphological remodeling of epithelial tissues. This suggests that keratin loci can be developed as sensitive biomarkers to assay temporal disruptions of larval-to-adult gene expression programs. Our study has identified the first collection of loci that are regulated during TH-induced metamorphosis in a salamander, thus setting the stage for future investigations of TH disruption in the Mexican axolotl and other salamanders of the genus Ambystoma.
Phylogenetic analysis of Common Garter Snake (Thamnophis sirtalis) stomach contents detects cryptic range of a secretive salamander (Ensatina eschscholtzii oregonensis) Herpetological Conservation and Biology 5(3):395–402
Sean B. Reilly; Andrew D Gottsho; Justin M. Garwood; Bryan. Jennings
Given the current global amphibian decline, it is crucial to obtain accurate and current information regarding species distributions. Secretive amphibians such as plethodontid salamanders can be difficult to detect in many cases, especially in remote, high elevation areas. We used molecular phylogenetic analyses to identify three partially digested salamanders palped...
Full Text Available A new fossil salamander, Nuominerpeton aquilonaris (gen. et sp. nov., is named and described based on specimens from the Lower Cretaceous Guanghua Formation of Inner Mongolia, China. The new discovery documents a far northern occurrence of Early Cretaceous salamanders in China, extending the geographic distribution for the Mesozoic fossil record of the group from the Jehol area (40th–45th parallel north to near the 49th parallel north. The new salamander is characterized by having the orbitosphenoid semicircular in shape; coracoid plate of the scapulocoracoid greatly expanded with a convex ventral and posterior border; ossification of two centralia in carpus and tarsus; and first digit being about half the length of the second digit in both manus and pes. The new salamander appears to be closely related to hynobiids, although this inferred relationship awaits confirmation by research in progress by us on a morphological and molecular combined analysis of cryptobranchoid relationships. Comparison of adult with larval and postmetamorphic juvenile specimens provides insights into developmental patterns of cranial and postcranial skeletons in this fossil species, especially resorption of the palatine and anterior portions of the palatopterygoid in the palate and the coronoid in the mandible during metamorphosis, and postmetamorphic ossification of the mesopodium in both manus and pes. Thus, this study provides a rare case study of developmental features in a Mesozoic salamander.
Perry, M.C.; Osenton, P.C.; Stoll, C.S.; Therres, Glenn D.
Amphibians, reptiles, birds, and mammals were surveyed at six created forested wetlands in central Maryland and at six adjacent reference forested wetlands during 1993-1996 to determine comparative biological diversity of these habitats. Amphibians and reptiles were caught in pitfall and funnel traps associated with 15.4m (50 ft) drift fences. Birds were surveyed with a complete count while walking through each area. Mammals were surveyed by capture in live traps. More species and total individuals of amphibians were caught on the reference wetlands than on the created wetlands. The red-backed salamander (Plethodon cinereus), the four-toed salamander (Hemidactylium scutatum), the eastern spadefoot (Scaphiopus holbrooki), and the wood frog (Rana sylvatica) were captured on the reference wetlands, but not on the created sites. The wood frog was captured at all reference sites and may represent the best amphibian species to characterize a forested wetland. Reptiles were not caught in sufficient numbers to warrant comparisons. Ninety-two bird species were recorded on created sites and 55 bird species on the reference sites. Bird species on the created sites represented those typically found in nonforested habitats. Mammal species were similar on both sites, but overall the reference sites had three times the number caught on created sites. The meadow vole (Microtus pennsylvanicus) was the dominant species captured on created sites, and the white-footed mouse (Peromyscus leucopus) was the dominant species on reference sites, with little habitat overlap for these two species. Although species richness and total number of animals were high for created forested wetlands, these survey results show major differences from species expected for a forested wetland. The created forested wetlands appear to provide good habitat for wildlife, but are probably not providing the full functions and values of the forested wetlands that they were constructed to replace.
Anderson, Thomas L; Ousterhout, Brittany H; Peterman, William E; Drake, Dana L; Semlitsch, Raymond D
Drought is a strong density-independent environmental filter that contributes to population regulation and other ecological processes. Not all species respond similarly to drought, and the overall impacts can vary depending on life histories. Such differences can necessitate management strategies that incorporate information on individual species to maximize conservation success. We report the effects of a short-term drought on occupancy and reproductive success of two pond-breeding salamanders that differ in breeding phenology (fall vs. spring breeder) across an active military base landscape in Missouri, USA: We surveyed ~200 ponds for the presence of eggs, larvae, and metamorphs from 2011 to 2013. This period coincided with before, during, and after a severe drought that occurred in 2012. The two species showed contrasting responses to drought, where high reproductive failure (34% of ponds) was observed for the spring breeder during a single drought year. Alternatively, the fall breeder only showed a cumulative 8% failure over two years. The number of breeding ponds available for use in the fall decreased during the drought due to pond drying and/or a lack of re-filling. Estimates of occupancy probability declined for the fall-breeding salamander between 2012 and 2013, whereas occupancy probability estimates of the spring breeder increased post-drought. The presence of fish, hydroperiod, the amount of forest cover surrounding ponds, and canopy cover were all found to affect estimates of occupancy probabilities of each species. Pond clustering (distance to nearest pond and the number of ponds within close proximity), hydroperiod, forest cover, and canopy cover influenced both estimates of colonization and extinction probabilities. Our results show life history variation can be important in determining the relative susceptibility of a species to drought conditions, and that sympatric species experiencing the same environmental conditions can respond differently
Bendik, Nathan F; Meik, Jesse M; Gluesenkamp, Andrew G; Roelke, Corey E; Chippindale, Paul T
Subterranean faunal radiations can result in complex patterns of morphological divergence involving both convergent or parallel phenotypic evolution and cryptic species diversity. Salamanders of the genus Eurycea in central Texas provide a particularly challenging example with respect to phylogeny reconstruction, biogeography and taxonomy. These predominantly aquatic species inhabit karst limestone aquifers and spring outflows, and exhibit a wide range of morphological and genetic variation. We extensively sampled spring and cave populations of six Eurycea species within this group (eastern Blepsimolge clade), to reconstruct their phylogenetic and biogeographic history using mtDNA and examine patterns and origins of cave- and surface-associated morphological variation. Genetic divergence is generally low, and many populations share ancestral haplotypes and/or show evidence of introgression. This pattern likely indicates a recent radiation coupled with a complex history of intermittent connections within the aquatic karst system. Cave populations that exhibit the most extreme troglobitic morphologies show no or very low divergence from surface populations and are geographically interspersed among them, suggesting multiple instances of rapid, parallel phenotypic evolution. Morphological variation is diffuse among cave populations; this is in contrast to surface populations, which form a tight cluster in morphospace. Unexpectedly, our analyses reveal two distinct and previously unrecognized morphological groups encompassing multiple species that are not correlated with spring or cave habitat, phylogeny or geography, and may be due to developmental plasticity. The evolutionary history of this group of spring- and cave-dwelling salamanders reflects patterns of intermittent isolation and gene flow influenced by complex hydrogeologic dynamics that are characteristic of karst regions. Shallow genetic divergences among several species, evidence of genetic exchange, and
Rovito Sean M
Full Text Available Abstract Background The complex geological history of Mesoamerica provides the opportunity to study the impact of multiple biogeographic barriers on population differentiation. We examine phylogeographic patterns in a clade of lowland salamanders (Bolitoglossa subgenus Nanotriton using two mitochondrial genes and one nuclear gene. We use several phylogeographic analyses to infer the history of this clade and test hypotheses regarding the geographic origin of species and location of genetic breaks within species. We compare our results to those for other taxa to determine if historical events impacted different species in a similar manner. Results Deep genetic divergence between species indicates that they are relatively old, and two of the three widespread species show strong phylogeographic structure. Comparison of mtDNA and nuclear gene trees shows no evidence of hybridization or introgression between species. Isolated populations of Bolitoglossa rufescens from Los Tuxtlas region constitute a separate lineage based on molecular data and morphology, and divergence between Los Tuxtlas and other areas appears to predate the arrival of B. rufescens in other areas west of the Isthmus of Tehuantepec. The Isthmus appears responsible for Pliocene vicariance within B. rufescens, as has been shown for other taxa. The Motagua-Polochic fault system does not appear to have caused population vicariance, unlike in other systems. Conclusions Species of Nanotriton have responded to some major geological events in the same manner as other taxa, particularly in the case of the Isthmus of Tehuantepec. The deep divergence of the Los Tuxtlas populations of B. rufescens from other populations highlights the contribution of this volcanic system to patterns of regional endemism, and morphological differences observed in the Los Tuxtlas populations suggests that they may represent an undescribed species of Bolitoglossa. The absence of phylogeographic structure in B
Full Text Available Next-generation technologies for determination of genomics and transcriptomics composition have a wide range of applications. Andrias davidianus, has become an endangered amphibian species of salamander endemic in China. However, there is a lack of the molecular information. In this study, we obtained the RNA-Seq data from a pool of A. davidianus tissue including spleen, liver, muscle, kidney, skin, testis, gut and heart using Illumina HiSeq 2500 platform. A total of 15,398,997,600 bp were obtained, corresponding to 102,659,984 raw reads. A total of 102,659,984 reads were filtered after removing low-quality reads and trimming the adapter sequences. The Trinity program was used to de novo assemble 132,912 unigenes with an average length of 690 bp and N50 of 1263 bp. Unigenes were annotated through number of databases. These transcriptomic data of A. davidianus should open the door to molecular evolution studies based on the entire transcriptome or targeted genes of interest to sequence. The raw data in this study can be available in NCBI SRA database with accession number of SRP099564.
Yoshikawa, Natsuhiko; Matsui, Masafumi
We describe two new species of hynobiid salamanders in the genus Onychodactylus from eastern Honshu, Japan, based on the morphological and genetic evidence. Onychodactylus intermedius sp. nov. is distributed in southern part of Tohoku District and northern Ibaraki and Niigata Prefectures, and was previously reported as S-Tohoku group. Onychodactylus intermedius belongs to the O. japonicus species complex, and differs from the other congeners in having relatively long tail, narrow head, short snout, 18 presacral vertebrae, and distinctly curved vomerine tooth series without gap. Onychodactylus fuscus sp. nov. is known from only four localities in Fukushima and Niigata Prefectures of Tohoku and Hokuriku Districts. It also belongs to the O. japonicus complex, but lacks the dorsal stripe, which is a diagnostic character of the species complex. In other characteristics, O. fuscus differs from the other congeners in having comparatively long tail, wide head and internarial space, shallowly curved vomerine tooth series with gap, and relatively few vomerine teeth. Both species described here breed in winter. Phylogenetically, the two new species are closely related to each other, forming a well-supported clade with O. tsukubaensis as their sister species. Onychodactylus intermedius sp. nov. is known to be parapatric with O. japonicus and O. nipponoborealis without hybridization, whereas O. fuscus sp. nov. is sympatric with O. japonicus at least in a single known locality, and analysis of microsatellite loci indicates they are reproductively isolated.
Full Text Available North America and the neotropics harbor nearly all species of plethodontid salamanders. In contrast, this family of caudate amphibians is represented in Europe and Asia by two genera, Speleomantes and Karsenia, which are confined to small geographic ranges. Compared to neotropical and North American plethodontids, mortality attributed to chytridiomycosis caused by Batrachochytrium dendrobatidis (Bd has not been reported for European plethodontids, despite the established presence of Bd in their geographic distribution. We determined the extent to which Bd is present in populations of all eight species of European Speleomantes and show that Bd was undetectable in 921 skin swabs. We then compared the susceptibility of one of these species, Speleomantes strinatii, to experimental infection with a highly virulent isolate of Bd (BdGPL, and compared this to the susceptible species Alytes muletensis. Whereas the inoculated A. muletensis developed increasing Bd-loads over a 4-week period, none of five exposed S. strinatii were colonized by Bd beyond 2 weeks post inoculation. Finally, we determined the extent to which skin secretions of Speleomantes species are capable of killing Bd. Skin secretions of seven Speleomantes species showed pronounced killing activity against Bd over 24 hours. In conclusion, the absence of Bd in Speleomantes combined with resistance to experimental chytridiomycosis and highly efficient skin defenses indicate that the genus Speleomantes is a taxon unlikely to decline due to Bd.
Rachael Y Dudaniec
Full Text Available With predicted decreases in genetic diversity and greater genetic differentiation at range peripheries relative to their cores, it can be difficult to distinguish between the roles of current disturbance versus historic processes in shaping contemporary genetic patterns. To address this problem, we test for differences in historic demography and landscape genetic structure of coastal giant salamanders (Dicamptodon tenebrosus in two core regions (Washington State, United States versus the species' northern peripheral region (British Columbia, Canada where the species is listed as threatened. Coalescent-based demographic simulations were consistent with a pattern of post-glacial range expansion, with both ancestral and current estimates of effective population size being much larger within the core region relative to the periphery. However, contrary to predictions of recent human-induced population decline in the less genetically diverse peripheral region, there was no genetic signature of population size change. Effects of current demographic processes on genetic structure were evident using a resistance-based landscape genetics approach. Among core populations, genetic structure was best explained by length of the growing season and isolation by resistance (i.e. a 'flat' landscape, but at the periphery, topography (slope and elevation had the greatest influence on genetic structure. Although reduced genetic variation at the range periphery of D. tenebrosus appears to be largely the result of biogeographical history rather than recent impacts, our analyses suggest that inherent landscape features act to alter dispersal pathways uniquely in different parts of the species' geographic range, with implications for habitat management.
Canestrelli, Daniele; Bisconti, Roberta; Nascetti, Giuseppe
Hybridization and introgression, contrary to previous beliefs, are now considered to be widespread processes even among animal species. Nonetheless, the range of their possible outcomes and roles in moulding biodiversity patterns are still far from being fully appraised. Here we investigated the pattern of hybridization and introgression between Salamandrina perspicillata and S. terdigitata, two salamanders endemic to the Italian peninsula. Using a set of diagnostic or differentiated genetic markers (9 nuclear and 1 mitochondrial), we documented extensive unidirectional introgression of S. terdigitata alleles into the S. perspicillata gene pool in central Italy, indicating that barriers against hybridization were permeable when they came into secondary contact, and despite their ancient divergence. Nonetheless, purebred S. terdigitata, as well as F1, F2, and backcrosses were not found within the hybrid zone. Moreover, Bayesian analyses of population structure identified admixed populations belonging to a differentiated gene pool with respect to both parental populations. Overall, the observed genetic structure, together with their geographic pattern of distribution, suggests that Salamandrina populations in central Italy could have entered a distinct evolutionary pathway. How far they have gone along this pathway will deserve future investigation. PMID:25269625
Rovito, Sean M.; García-París, Mario; Maisano, Jessica A.; Wake, David B.
We describe three new species of minute salamanders, genus Thorius, from the Sierra Madre del Sur of Oaxaca, Mexico. Until now only a single species, T. minutissimus, has been reported from this region, although molecular data have long shown extensive genetic differentiation among geographically disjunct populations. Adult Thorius pinicola sp. nov., T. longicaudus sp. nov., and T. tlaxiacus sp. nov. are larger than T. minutissimus and possess elliptical rather than oval nostrils; T. pinicola and T. longicaudus also have longer tails. All three new species occur west of the range of T. minutissimus, which has the easternmost distribution of any member of the genus. The new species are distinguished from each other and from other named Thorius in Oaxaca by a combination of adult body size, external morphology and osteology, and by protein characters (allozymes) and differences in DNA sequences. In addition, we redescribe T. minutissimus and a related species, T. narisovalis, to further clarify the taxonomic status of Oaxacan populations and to facilitate future studies of the remaining genetically differentiated Thorius that cannot be satisfactorily assigned to any named species. Populations of all five species considered here appear to have declined dramatically over the last one or two decades and live specimens are difficult to find in nature. Thorius may be the most endangered genus of amphibians in the world. All species may go extinct before the end of this century. PMID:27896029
Noah D Charney
Full Text Available Improving detection rates for elusive species with clumped distributions is often accomplished through adaptive sampling designs. This approach can be extended to include species with temporally variable detection probabilities. By concentrating survey effort in years when the focal species are most abundant or visible, overall detection rates can be improved. This requires either long-term monitoring at a few locations where the species are known to occur or models capable of predicting population trends using climatic and demographic data. For marbled salamanders (Ambystoma opacum in Massachusetts, we demonstrate that annual variation in detection probability of larvae is regionally correlated. In our data, the difference in survey success between years was far more important than the difference among the three survey methods we employed: diurnal surveys, nocturnal surveys, and dipnet surveys. Based on these data, we simulate future surveys to locate unknown populations under a temporally adaptive sampling framework. In the simulations, when pond dynamics are correlated over the focal region, the temporally adaptive design improved mean survey success by as much as 26% over a non-adaptive sampling design. Employing a temporally adaptive strategy costs very little, is simple, and has the potential to substantially improve the efficient use of scarce conservation funds.
Full Text Available We describe three new species of minute salamanders, genus Thorius, from the Sierra Madre del Sur of Oaxaca, Mexico. Until now only a single species, T. minutissimus, has been reported from this region, although molecular data have long shown extensive genetic differentiation among geographically disjunct populations. Adult Thorius pinicola sp. nov., T. longicaudus sp. nov., and T. tlaxiacus sp. nov. are larger than T. minutissimus and possess elliptical rather than oval nostrils; T. pinicola and T. longicaudus also have longer tails. All three new species occur west of the range of T. minutissimus, which has the easternmost distribution of any member of the genus. The new species are distinguished from each other and from other named Thorius in Oaxaca by a combination of adult body size, external morphology and osteology, and by protein characters (allozymes and differences in DNA sequences. In addition, we redescribe T. minutissimus and a related species, T. narisovalis, to further clarify the taxonomic status of Oaxacan populations and to facilitate future studies of the remaining genetically differentiated Thorius that cannot be satisfactorily assigned to any named species. Populations of all five species considered here appear to have declined dramatically over the last one or two decades and live specimens are difficult to find in nature. Thorius may be the most endangered genus of amphibians in the world. All species may go extinct before the end of this century.
Pasmans, Frank; Van Rooij, Pascale; Blooi, Mark; Tessa, Giulia; Bogaerts, Sergé; Sotgiu, Giuseppe; Garner, Trenton W J; Fisher, Matthew C; Schmidt, Benedikt R; Woeltjes, Tonnie; Beukema, Wouter; Bovero, Stefano; Adriaensen, Connie; Oneto, Fabrizio; Ottonello, Dario; Martel, An; Salvidio, Sebastiano
North America and the neotropics harbor nearly all species of plethodontid salamanders. In contrast, this family of caudate amphibians is represented in Europe and Asia by two genera, Speleomantes and Karsenia, which are confined to small geographic ranges. Compared to neotropical and North American plethodontids, mortality attributed to chytridiomycosis caused by Batrachochytrium dendrobatidis (Bd) has not been reported for European plethodontids, despite the established presence of Bd in their geographic distribution. We determined the extent to which Bd is present in populations of all eight species of European Speleomantes and show that Bd was undetectable in 921 skin swabs. We then compared the susceptibility of one of these species, Speleomantes strinatii, to experimental infection with a highly virulent isolate of Bd (BdGPL), and compared this to the susceptible species Alytes muletensis. Whereas the inoculated A. muletensis developed increasing Bd-loads over a 4-week period, none of five exposed S. strinatii were colonized by Bd beyond 2 weeks post inoculation. Finally, we determined the extent to which skin secretions of Speleomantes species are capable of killing Bd. Skin secretions of seven Speleomantes species showed pronounced killing activity against Bd over 24 hours. In conclusion, the absence of Bd in Speleomantes combined with resistance to experimental chytridiomycosis and highly efficient skin defenses indicate that the genus Speleomantes is a taxon unlikely to decline due to Bd.
Semiltsch, Raymond D; Walls, Susan; Barichivich, William J.; O'Donnell, Katherine
A comprehensive view of population declines and their underlying causes is necessary to reverse species loss. Historically, in many cases, a narrow view may have allowed species declines to continue, virtually undetected, for long periods of time (perhaps even decades). We suggest that extinction debt is likely responsible for numerous (perhaps most) amphibian declines and that this perspective should be incorporated into the structure of amphibian research and management. Extinction debt, originally proposed to explain changes in species richness following environmental disturbance, also may refer to the proportion of populations of an individual species that is expected to eventually be lost because of habitat change. A conservation framework to address extinction debt focuses research on threats at the individual, population, and metapopulation levels. This approach will help enhance, restore, and protect specific processes and habitats at the proper scale by directing management to the most vulnerable level and stage of a species. We illustrate this approach using Flatwoods Salamanders, Ambystoma cingulatumand Ambystoma bishopi, which occurred historically throughout the Coastal Plain of the southeastern United States but have experienced a greater than 85% loss of populations in recent years. Reversal of these losses is possible only if conservation and recovery efforts encompass individual, population, and metapopulation levels. We illustrate our framework by outlining actions that could be taken at each of these levels to help guide conservation and management of amphibians with complex life cycles and provide options for how to prioritize conservation actions in the face of logistical and budgetary shortfalls.
Balogová, Monika; Nelson, Emma; Uhrin, Marcel; Figurová, Mária; Ledecký, Valent; Zyśk, Bartłomiej
It has been proposed that digit ratio may be used as a biomarker of early developmental effects. Specifically, the second-to-fourth digit ratio (2D:4D) has been linked to the effects of sex hormones and their receptor genes, but other digit ratios have also been investigated. Across taxa, patterns of sexual dimorphism in digit ratios are ambiguous and a scarcity of studies in basal tetrapods makes it difficult to understand how ratios have evolved. Here, we focus on examining sex differences in digit ratios (2D:3D, 2D:4D, and 3D:4D) in a common amphibian, the fire salamander (Salamandra salamandra). We used graphic software to measure soft tissue digit length and digit bone length from X-rays. We found a nonsignificant tendency in males to have a lower 2D:3D than females; however, no sexual differences were detected in the other ratios. We discuss our results in the context of other studies of digit ratios, and how sex determination systems, as well as other factors, might impact patterns of sexual dimorphism, particularly in reptiles and in amphibians. Our findings suggest that caution is needed when using digit ratios as a potential indicator of prenatal hormonal effects in amphibians and highlight the need for more comparative studies to elucidate the evolutionary and genetic mechanisms implicated in sexually dimorphic patterns across taxonomic groups. © 2015 Wiley Periodicals, Inc.
Cosentino, Bradley J; Phillips, Christopher A; Schooley, Robert L; Lowe, Winsor H; Douglas, Marlis R
Theory predicts that founder effects have a primary role in determining metapopulation genetic structure. However, ecological factors that affect extinction-colonization dynamics may also create spatial variation in the strength of genetic drift and migration. We tested the hypothesis that ecological factors underlying extinction-colonization dynamics influenced the genetic structure of a tiger salamander (Ambystoma tigrinum) metapopulation. We used empirical data on metapopulation dynamics to make a priori predictions about the effects of population age and ecological factors on genetic diversity and divergence among 41 populations. Metapopulation dynamics of A. tigrinum depended on wetland area, connectivity and presence of predatory fish. We found that newly colonized populations were more genetically differentiated than established populations, suggesting that founder effects influenced genetic structure. However, ecological drivers of metapopulation dynamics were more important than age in predicting genetic structure. Consistent with demographic predictions from metapopulation theory, genetic diversity and divergence depended on wetland area and connectivity. Divergence was greatest in small, isolated wetlands where genetic diversity was low. Our results show that ecological factors underlying metapopulation dynamics can be key determinants of spatial genetic structure, and that habitat area and isolation may mediate the contributions of drift and migration to divergence and evolution in local populations.
Zhang, Hui; Zhong, Shengwei; Yu, Pengcheng; Ge, Tingting; Peng, Shasha; Guo, Xiaoquan; Zhou, Zuohong
In this study, we attempt to identify gastric telocytes (TCs) of the Chinese giant salamander Andrias davidianus, by light microscopy, immunohistochemistry and transmission electron microscopy (TEM) methods. Toluidine blue staining showed TCs with one to two very thin and long telopodes (Tps) that were located in gastric lamina propria. Tps had characteristic structures, including podoms, podomers and dichotomous branching. Immunohistochemistry showed the existence of CD34+/PDGFRα+ TCs with moniliform Tps in stroma and were close to gastric glands and blood vessels. TEM micrographs also demonstrated the presence of TCs in interstitium between gastric glands. TCs/Tps were located in close proximity to gastric glands, blood vessels, endocrine cells and stem cells. In particular, Tps frequently surrounded stem cells. TCs and Tps, Tps and stem cells established close contacts. Moreover, the exosomes were also found near TCs/Tps. Our data confirmed the presence of TCs in gastric lamina propria of the amphibian, and suggested that TCs cooperate with resident stem cells to regulate endocrine cells and gastric glands regeneration and homeostasis. PMID:27629815
Isayama, Tomoki; McCabe England, S L; Crouch, R K; Zimmerman, A L; Makino, C L
Vision begins with photoisomerization of 11-cis retinal to the all-trans conformation within the chromophore-binding pocket of opsin, leading to activation of a biochemical cascade. Release of all-trans retinal from the binding pocket curtails but does not fully quench the ability of opsin to activate transducin. All-trans retinal and some other analogs, such as beta-ionone, enhance opsin's activity, presumably on binding the empty chromophore-binding pocket. By recording from isolated salamander photoreceptors and from patches of rod outer segment membrane, we now show that high concentrations of beta-ionone suppressed circulating current in dark-adapted green-sensitive rods by inhibiting the cyclic nucleotide-gated channels. There were also decreases in circulating current and flash sensitivity, and accelerated flash response kinetics in dark-adapted blue-sensitive (BS) rods and cones, and in ultraviolet-sensitive cones, at concentrations too low to inhibit the channels. These effects persisted in BS rods even after incubation with 9-cis retinal to ensure complete regeneration of their visual pigment. After long exposures to high concentrations of beta-ionone, recovery was incomplete unless 9-cis retinal was given, indicating that visual pigment had been bleached. Therefore, we propose that beta-ionone activates and bleaches some types of visual pigments, mimicking the effects of light.
McCusker, Catherine D; Gardiner, David M
Regenerative medicine has reached the point where we are performing clinical trials with stem-cell-derived cell populations in an effort to treat numerous human pathologies. However, many of these efforts have been challenged by the inability of the engrafted populations to properly integrate into the host environment to make a functional biological unit. It is apparent that we must understand the basic biology of tissue integration in order to apply these principles to the development of regenerative therapies in humans. Studying tissue integration in model organisms, where the process of integration between the newly regenerated tissues and the 'old' existing structures can be observed and manipulated, can provide valuable insights. Embryonic and adult cells have a memory of their original position, and this positional information can modify surrounding tissues and drive the formation of new structures. In this Review, we discuss the positional interactions that control the ability of grafted cells to integrate into existing tissues during the process of salamander limb regeneration, and discuss how these insights could explain the integration defects observed in current cell-based regenerative therapies. Additionally, we describe potential molecular tools that can be used to manipulate the positional information in grafted cell populations, and to promote the communication of positional cues in the host environment to facilitate the integration of engrafted cells. Lastly, we explain how studying positional information in current cell-based therapies and in regenerating limbs could provide key insights to improve the integration of cell-based regenerative therapies in the future.
Savage, Wesley K; Fremier, Alexander K; Shaffer, H Bradley
Quantifying the influence of the landscape on the genetic structure of natural populations remains an important empirical challenge, particularly for poorly studied, ecologically cryptic species. We conducted an extensive microsatellite analysis to examine the population genetics of the southern long-toed salamander (Ambystoma macrodactylum sigillatum) in a naturally complex landscape. Using spatially explicit modelling, we investigated the influence of the Sierra Nevada topography on potential dispersal corridors between sampled populations. Our results indicate very high-genetic divergence among populations, high within-deme relatedness, and little evidence of recent migration or population admixture. We also discovered unexpectedly high between-year genetic differentiation (F(ST)) for breeding sites, suggesting that breeding groups vary over localized space and time. While environmental factors associated with high-elevation montane habitats apparently play an important role in shaping population differentiation, additional, species-specific biological processes must also be operating to account for observed deviations from temporal, among-year panmixia. Our study emphasizes the population-level insights that can be gained from high-density sampling in space and time, and the highly substructured population biology that may characterize amphibians in extreme montane habitats.
Mueller, Rachel Lockridge; Macey, J. Robert; Jaekel, Martin; Wake, David B.; Boore, Jeffrey L.
The evolutionary history of the largest salamander family (Plethodontidae) is characterized by extreme morphological homoplasy. Analysis of the mechanisms generating such homoplasy requires an independent, molecular phylogeny. To this end, we sequenced 24 complete mitochondrial genomes (22 plethodontids and two outgroup taxa), added data for three species from GenBank, and performed partitioned and unpartitioned Bayesian, ML, and MP phylogenetic analyses. We explored four dataset partitioning strategies to account for evolutionary process heterogeneity among genes and codon positions, all of which yielded increased model likelihoods and decreased numbers of supported nodes in the topologies (PP > 0.95) relative to the unpartitioned analysis. Our phylogenetic analyses yielded congruent trees that contrast with the traditional morphology-based taxonomy; the monophyly of three out of four major groups is rejected. Reanalysis of current hypotheses in light of these new evolutionary relationships suggests that (1) a larval life history stage re-evolved from a direct-developing ancestor multiple times, (2) there is no phylogenetic support for the ''Out of Appalachia'' hypothesis of plethodontid origins, and (3) novel scenarios must be reconstructed for the convergent evolution of projectile tongues, reduction in toe number, and specialization for defensive tail loss. Some of these novel scenarios imply morphological transformation series that proceed in the opposite direction than was previously thought. In addition, they suggest surprising evolutionary lability in traits previously interpreted to be conservative.
Charney, Noah D; Kubel, Jacob E; Eiseman, Charles S
Improving detection rates for elusive species with clumped distributions is often accomplished through adaptive sampling designs. This approach can be extended to include species with temporally variable detection probabilities. By concentrating survey effort in years when the focal species are most abundant or visible, overall detection rates can be improved. This requires either long-term monitoring at a few locations where the species are known to occur or models capable of predicting population trends using climatic and demographic data. For marbled salamanders (Ambystoma opacum) in Massachusetts, we demonstrate that annual variation in detection probability of larvae is regionally correlated. In our data, the difference in survey success between years was far more important than the difference among the three survey methods we employed: diurnal surveys, nocturnal surveys, and dipnet surveys. Based on these data, we simulate future surveys to locate unknown populations under a temporally adaptive sampling framework. In the simulations, when pond dynamics are correlated over the focal region, the temporally adaptive design improved mean survey success by as much as 26% over a non-adaptive sampling design. Employing a temporally adaptive strategy costs very little, is simple, and has the potential to substantially improve the efficient use of scarce conservation funds.
Anderson, Thomas L; Whiteman, Howard H
Assessment of the relative strengths of intra- and interspecific competition has increased in recent years and is critical to understanding the importance of competition. Yet, whether intra- and interspecific competition can have non-additive effects has rarely been tested. The resulting fitness consequences of such non-additive interactions are important to provide the context necessary to advance our understanding of competition theory. We compared the strength of additive and non-additive intra- and interspecific competition by manipulating densities of a pair of larval salamanders (Ambystoma talpoideum and A. maculatum) in experimental mesocosms within a response surface design. Intraspecific density had the strongest effect on the strength of competition for both species, and few observed comparisons indicated interspecific competition was an important factor in predicting body size, growth or larval period length of either species. Non-additive effects of intra- and interspecific competition influenced some response variables, including size and mass at metamorphosis in A. maculatum, but at a reduced strength compared to intraspecific effects alone. Intraspecific competition was thus the dominant biotic interaction, but non-additive effects also impact the outcome of competition in these species, validating the importance of testing for and incorporating non-additive density effects into competition models.
Aoki, Gen; Matsui, Masafumi; Nishikawa, Kanto
The Tohoku salamander, Hynobius lichenatus Boulenger, 1883, is a lentic breeding species widespread throughout montane regions of northeastern Japan. To explore intraspecific genetic variation and infer evolutionary history of H. lichenatus, we performed mitochondrial DNA analysis (complete 1141 bp sequences of the mitochondrial cytochrome b gene) using 215 adult and larval individuals collected from 75 localities, encompassing known distributional range of the species. Hynobius lichenatus proved to be monophyletic, including three well-supported and geographically structured clades (Clade I from northern Kanto, Clade II from southern Tohoku, and Clade III from northern Tohoku). These clades, respectively, comprise several subclades, and show genetic distances as large as those seen between different species of Hynobius. Results of population statistic analyses indicate that all clades and most subclades have maintained high genetic diversity and demographic stability over long periods. Molecular dating indicates divergence in H. lichenatus concords with topographic evolution of northeastern Japan from late Miocene to early Pleistocene, suggesting that paleogeographic events in this region, such as orogenesis, sea level change, and volcanic activity, have been crucial for shaping genetic patterns and diversity in this species. Hynobius lichenatus greatly differs from many other animal species from northeastern Japan in its much older periods and the pattern of genetic differentiation, and is suggested as an old faunal element in this region.
Full Text Available Abstract Background Amphibians are sensitive indicators of environmental conditions and show measurable responses, such as changes in phenology, abundance and range limits to local changes in precipitation and temperature regimes. Amphibians offer unique opportunities to study the important ecological and evolutionary implications of responses in life history characteristics to climatic change. We analyzed a late-Holocene fossil record of the Tiger Salamander (Ambystoma tigrinum for evidence of population-level changes in body size and paedomorphosis to climatic change over the last 3000 years. Results We found a significant difference in body size index between paedomorphic and metamorphic individuals during the time interval dominated by the Medieval Warm Period. There is a consistent ratio of paedomorphic to metamorphic specimens through the entire 3000 years, demonstrating that not all life history characteristics of the population were significantly altered by changes in climate on this timescale. Conclusion The fossil record of Ambystoma tigrinum we used spans an ecologically relevant timescale appropriate for understanding population and community response to projected climatic change. The population-level responses we documented are concordant with expectations based on modern environmental studies, and yield insight into population-level patterns across hundreds of generations, especially the independence of different life history characteristics. These conclusions lead us to offer general predictions about the future response of this species based on likely scenarios of climatic warming in the Rocky Mountain region.
Bonett, Ronald M; Chippindale, Paul T
Habitat variation strongly influences the evolution of developmentally flexible traits, and may drive speciation and diversification. The plethodontid salamander Eurycea tynerensis is endemic to the geologically diverse Ozark Plateau of south-central North America, and comprises both strictly aquatic paedomorphic populations (achieving reproductive maturity while remaining in the larval form) and more terrestrial metamorphic populations. The switch between developmental modes has occurred many times, but populations typically exhibit a single life history mode. This unique system offers an opportunity to study the specific ecological circumstances under which alternate developmental and life history modes evolve. We use phylogenetic independent contrasts to test for relationships between a key microhabitat feature (streambed sediment) and this major life history polymorphism. We find streambed microstructure (sediment particle size, type and degree of sorting) to be highly correlated with life-history mode. Eurycea tynerensis is paedomorphic in streams containing large chert gravel, but metamorphoses in nearby streams containing poorly sorted, clastic material such as sandstone or siltstone. Deposits of large chert gravel create loosely associated streambeds, which provide access to subsurface water during dry summer months. Conversely, streambeds composed of more densely packed sandstone and siltstone sediments leave no subterranean refuge when surface water dries, presumably necessitating metamorphosis and use of terrestrial habitats. This represents a clear example of the relationship between microhabitat structure and evolution of a major developmental and life history trait, and has broad implications for the role of localized ecological conditions on larger-scale evolutionary processes.
Xu, Jian Wei; Slaughter, Malcolm M
Large-conductance calcium-activated potassium (BK) channels are colocalized with calcium channels at sites of exocytosis at the presynaptic terminals throughout the nervous system. It is expected that their activation would provide negative feedback to transmitter release, but the opposite is sometimes observed. Attempts to resolve this apparent paradox based on alterations in action potential waveform have been ambiguous. In an alternative approach, we investigated the influence of this channel on neurotransmitter release in a nonspiking neuron, the salamander rod photoreceptors. Surprisingly, the BK channel facilitates calcium-mediated transmitter release from rods. The two presynaptic channels form a positive coupled loop. Calcium influx activates the BK channel current, leading to potassium efflux that increases the calcium current. The normal physiological voltage range of the rod is well matched to the dynamics of this positive loop. When the rod is further depolarized, then the hyperpolarizing BK channel current exceeds its facilitatory effect, causing truncation of transmitter release. Thus, the calcium channel-BK channel linkage performs two functions at the synapse: nonlinear potentiator and safety brake.
McMenamin, Sarah K; Hadly, Elizabeth A
Recent data indicates that blotched tiger salamanders (Ambystoma tigrinum melanostictum) in northern regions of Yellowstone National Park are declining due to climate-related habitat changes. In this study, we used ancient and modern mitochondrial haplotype diversity to model the effective size of this amphibian population through recent geological time and to assess past responses to climatic changes in the region. Using subfossils collected from a cave in northern Yellowstone, we analyzed >700 base pairs of mitochondrial sequence from 16 samples ranging in age from 100 to 3300 years old and found that all shared an identical haplotype. Although mitochondrial diversity was extremely low within the living population, we still were able to detect geographic subdivision within the local area. Using serial coalescent modelling with Bayesian priors from both modern and ancient genetic data we simulated a range of probable population sizes and mutation rates through time. Our simulations suggest that regional mitochondrial diversity has remained relatively constant even through climatic fluctuations of recent millennia.
Sarah K McMenamin
Full Text Available Recent data indicates that blotched tiger salamanders (Ambystoma tigrinum melanostictum in northern regions of Yellowstone National Park are declining due to climate-related habitat changes. In this study, we used ancient and modern mitochondrial haplotype diversity to model the effective size of this amphibian population through recent geological time and to assess past responses to climatic changes in the region. Using subfossils collected from a cave in northern Yellowstone, we analyzed >700 base pairs of mitochondrial sequence from 16 samples ranging in age from 100 to 3300 years old and found that all shared an identical haplotype. Although mitochondrial diversity was extremely low within the living population, we still were able to detect geographic subdivision within the local area. Using serial coalescent modelling with Bayesian priors from both modern and ancient genetic data we simulated a range of probable population sizes and mutation rates through time. Our simulations suggest that regional mitochondrial diversity has remained relatively constant even through climatic fluctuations of recent millennia.
Parra-Olea, Gabriela; Rovito, Sean M; García-París, Mario; Maisano, Jessica A; Wake, David B; Hanken, James
We describe three new species of minute salamanders, genus Thorius, from the Sierra Madre del Sur of Oaxaca, Mexico. Until now only a single species, T. minutissimus, has been reported from this region, although molecular data have long shown extensive genetic differentiation among geographically disjunct populations. Adult Thorius pinicola sp. nov., T. longicaudus sp. nov., and T. tlaxiacus sp. nov. are larger than T. minutissimus and possess elliptical rather than oval nostrils; T. pinicola and T. longicaudus also have longer tails. All three new species occur west of the range of T. minutissimus, which has the easternmost distribution of any member of the genus. The new species are distinguished from each other and from other named Thorius in Oaxaca by a combination of adult body size, external morphology and osteology, and by protein characters (allozymes) and differences in DNA sequences. In addition, we redescribe T. minutissimus and a related species, T. narisovalis, to further clarify the taxonomic status of Oaxacan populations and to facilitate future studies of the remaining genetically differentiated Thorius that cannot be satisfactorily assigned to any named species. Populations of all five species considered here appear to have declined dramatically over the last one or two decades and live specimens are difficult to find in nature. Thorius may be the most endangered genus of amphibians in the world. All species may go extinct before the end of this century.
Shaffer H Bradley
Full Text Available Abstract Background Hybrid zones represent valuable opportunities to observe evolution in systems that are unusually dynamic and where the potential for the origin of novelty and rapid adaptation co-occur with the potential for dysfunction. Recently initiated hybrid zones are particularly exciting evolutionary experiments because ongoing natural selection on novel genetic combinations can be studied in ecological time. Moreover, when hybrid zones involve native and introduced species, complex genetic patterns present important challenges for conservation policy. To assess variation of admixture dynamics, we scored a large panel of markers in five wild hybrid populations formed when Barred Tiger Salamanders were introduced into the range of California Tiger Salamanders. Results At three of 64 markers, introduced alleles have largely displaced native alleles within the hybrid populations. Another marker (GNAT1 showed consistent heterozygote deficits in the wild, and this marker was associated with embryonic mortality in laboratory F2's. Other deviations from equilibrium expectations were idiosyncratic among breeding ponds, consistent with highly stochastic demographic effects. Conclusion While most markers retain native and introduced alleles in expected proportions, strong selection appears to be eliminating native alleles at a smaller set of loci. Such rapid fixation of alleles is detectable only in recently formed hybrid zones, though it might be representative of dynamics that frequently occur in nature. These results underscore the variable and mosaic nature of hybrid genomes and illustrate the potency of recombination and selection in promoting variable, and often unpredictable genetic outcomes. Introgression of a few, strongly selected introduced alleles should not necessarily affect the conservation status of California Tiger Salamanders, but suggests that genetically pure populations of this endangered species will be difficult to
Shaffer H Bradley
Full Text Available Abstract Background Introductions of non-native tiger salamanders into the range of California tiger salamanders have provided a rare opportunity to study the early stages of secondary contact and hybridization. We produced first- and second-generation hybrid salamanders in the lab and measured viability among these early-generation hybrid crosses to determine the strength of the initial barrier to gene exchange. We also created contemporary-generation hybrids in the lab and evaluated the extent to which selection has affected fitness over approximately 20 generations of admixture. Additionally, we examined the inheritance of quantitative phenotypic variation to better understand how evolution has progressed since secondary contact. Results We found significant variation in the fitness of hybrids, with non-native backcrosses experiencing the highest survival and F2 hybrids the lowest. Contemporary-generation hybrids had similar survival to that of F2 families, contrary to our expectation that 20 generations of selection in the wild would eliminate unfit genotypes and increase survival. Hybrid survival clearly exhibited effects of epistasis, whereas size and growth showed mostly additive genetic variance, and time to metamorphosis showed substantial dominance. Conclusions Based on first- and second- generation cross types, our results suggest that the initial barrier to gene flow between these two species was relatively weak, and subsequent evolution has been generally slow. The persistence of low-viability recombinant hybrid genotypes in some contemporary populations illustrates that while hybridization can provide a potent source of genetic variation upon which natural selection can act, the sorting of fit from unfit gene combinations might be inefficient in highly admixed populations. Spatio-temporal fluctuation in selection or complex genetics has perhaps stalled adaptive evolution in this system despite selection for admixed genotypes within
Germain, Damien; Laurin, Michel
Ossification sequences of the skull in extant Urodela and in Permo-Carboniferous Branchiosauridae have already been used to study the origin of lissamphibians. But most of these studies did not consider some recent methods developed to analyze the developmental sequences within a phylogenetic framework. Here, we analyze the ossification sequences of 24 cranial bones of 23 extant species of salamanders using the event-pairing method. This reveals new developmental synapomorphies for several extant salamander taxa and ancestral sequences for Urodela under four alternative reference phylogenies. An analysis with the 12 bones for which ossification sequence data are available in urodeles and in the branchiosaurid Apateon is also performed in order to compare the ancestral condition of the crown-group of Urodela to the sequence of Apateon. This reveals far more incompatibilities than previously suggested. The similarities observed between some extant salamanders and branchiosaurids may result from extensive homoplasy, as the extreme variation observed in extant Urodela suggests, or be plesiomorphic, as the conservation of some ossification patterns observed in other remotely related vertebrates like actinopterygians suggests. We propose a new, simpler method based on squared-change optimization to estimate the relative timing of ossification of various bones of hypothetical ancestors, and use independent-contrasts analysis to estimate the confidence intervals around these times. Our results show that the uncertainty of the ancestral ossification sequence of Urodela is much greater than event-pairing suggests. The developmental data do not allow to conclude that branchiosaurids are closely related to salamanders and their limited taxonomic distribution in Paleozoic taxa precludes testing hypotheses about lissamphibian origins. This is true regardless of the analytical method used (event-pairing or our new method based on squared-change parsimony). Simulations show that
Krause, E.T.; Caspers, B.A.
Fire salamanders are amphibians that exhibit a highly specific reproductive mode termed ovo-viviparity. The eggs develop inside their mothers, and the females give birth to fully developed larvae. The larvae in our study area cluster in two distinct genetic groups that can be linked directly to the
Fortuny, Josep; Marcé-Nogué, Jordi; Heiss, Egon; Sanchez, Montserrat; Gil, Lluis; Galobart, Àngel
Biting is an integral feature of the feeding mechanism for aquatic and terrestrial salamanders to capture, fix or immobilize elusive or struggling prey. However, little information is available on how it works and the functional implications of this biting system in amphibians although such approaches might be essential to understand feeding systems performed by early tetrapods. Herein, the skull biomechanics of the Chinese giant salamander, Andrias davidianus is investigated using 3D finite element analysis. The results reveal that the prey contact position is crucial for the structural performance of the skull, which is probably related to the lack of a bony bridge between the posterior end of the maxilla and the anterior quadrato-squamosal region. Giant salamanders perform asymmetrical strikes. These strikes are unusual and specialized behavior but might indeed be beneficial in such sit-and-wait or ambush-predators to capture laterally approaching prey. However, once captured by an asymmetrical strike, large, elusive and struggling prey have to be brought to the anterior jaw region to be subdued by a strong bite. Given their basal position within extant salamanders and their "conservative" morphology, cryptobranchids may be useful models to reconstruct the feeding ecology and biomechanics of different members of early tetrapods and amphibians, with similar osteological and myological constraints.
Krause, E.T.; Caspers, B.A.
Fire salamanders are amphibians that exhibit a highly specific reproductive mode termed ovo-viviparity. The eggs develop inside their mothers, and the females give birth to fully developed larvae. The larvae in our study area cluster in two distinct genetic groups that can be linked directly to the
Full Text Available Biting is an integral feature of the feeding mechanism for aquatic and terrestrial salamanders to capture, fix or immobilize elusive or struggling prey. However, little information is available on how it works and the functional implications of this biting system in amphibians although such approaches might be essential to understand feeding systems performed by early tetrapods. Herein, the skull biomechanics of the Chinese giant salamander, Andrias davidianus is investigated using 3D finite element analysis. The results reveal that the prey contact position is crucial for the structural performance of the skull, which is probably related to the lack of a bony bridge between the posterior end of the maxilla and the anterior quadrato-squamosal region. Giant salamanders perform asymmetrical strikes. These strikes are unusual and specialized behavior but might indeed be beneficial in such sit-and-wait or ambush-predators to capture laterally approaching prey. However, once captured by an asymmetrical strike, large, elusive and struggling prey have to be brought to the anterior jaw region to be subdued by a strong bite. Given their basal position within extant salamanders and their "conservative" morphology, cryptobranchids may be useful models to reconstruct the feeding ecology and biomechanics of different members of early tetrapods and amphibians, with similar osteological and myological constraints.
Heiss, Egon; Natchev, Nikolay; Gumpenberger, Michaela; Weissenbacher, Anton; Van Wassenbergh, Sam
During the evolutionary transition from fish to tetrapods, a shift from uni- to bidirectional suction feeding systems followed a reduction in the gill apparatus. Such a shift can still be observed during metamorphosis of salamanders, although many adult salamanders retain their aquatic lifestyle and feed by high-performance suction. Unfortunately, little is known about the interplay between jaws and hyobranchial motions to generate bidirectional suction flows. Here, we study the cranial morphology, as well as kinematic and hydrodynamic aspects related to prey capture in the Chinese giant salamander (Andrias davidianus). Compared with fish and previously studied amphibians, A. davidianus uses an alternative suction mechanism that mainly relies on accelerating water by separating the ‘plates’ formed by the long and broad upper and lower jaw surfaces. Computational fluid dynamics simulations, based on three-dimensional morphology and kinematical data from high-speed videos, indicate that the viscerocranial elements mainly serve to accommodate the water that was given a sufficient anterior-to-posterior impulse beforehand by powerful jaw separation. We hypothesize that this modified way of generating suction is primitive for salamanders, and that this behaviour could have played an important role in the evolution of terrestrial life in vertebrates by releasing mechanical constraints on the hyobranchial system. PMID:23466557
Full Text Available Abstract Background A major goal in evolutionary biology is to understand the evolution of phenotypic diversity. Both natural and sexual selection play a large role in generating phenotypic adaptations, with biomechanical requirements and developmental mechanisms mediating patterns of phenotypic evolution. For many traits, the relative importance of selective and developmental components remains understudied. Results We investigated ontogenetic trajectories of foot morphology in the eight species of European plethodontid cave salamander to test the hypothesis that adult foot morphology was adapted for climbing. Using geometric morphometrics and other approaches, we found that developmental patterns in five species displayed little morphological change during growth (isometry, where the extensive interdigital webbing in adults was best explained as the retention of the juvenile morphological state. By contrast, three species exhibited significant allometry, with an increase in interdigital webbing during growth. Phylogenetic analyses revealed that multiple evolutionary transitions between isometry and allometry of foot webbing have occurred in this lineage. Allometric parameters of foot growth were most similar to those of a tropical species previously shown to be adapted for climbing. Finally, interspecific variation in adult foot morphology was significantly reduced as compared to variation among juveniles, indicating that ontogenetic convergence had resulted in a common adult foot morphology across species. Conclusions The results presented here provide evidence of a complex history of phenotypic evolution in this clade. The common adult phenotype exhibited among species reveals that selection plays an important part in generating patterns of foot diversity in the group. However, developmental trajectories arriving at this common morphology are distinct; with some species displaying developmental stasis (isometry, while others show an increase
Anderson, Thomas L; Hocking, Daniel J; Conner, Christopher A; Earl, Julia E; Harper, Elizabeth B; Osbourn, Michael S; Peterman, William E; Rittenhouse, Tracy A G; Semlitsch, Raymond D
Phenology often determines the outcome of interspecific interactions, where early-arriving species often dominate interactions over those arriving later. The effects of phenology on species interactions are especially pronounced in aquatic systems, but the evidence is largely derived from experimental studies. We examined whether differences in breeding phenology between two pond-breeding salamanders (Ambystoma annulatum and A. maculatum) affected metamorph recruitment and demographic traits within natural populations, with the expectation that the fall-breeding A. annulatum would negatively affect the spring-breeding A. maculatum. We monitored populations of each species at five ponds over 4 years using drift fences. Metamorph abundance and survival of A. annulatum were affected by intra- and interspecific processes, whereas metamorph size and date of emigration were primarily influenced by intraspecific effects. Metamorph abundance, snout-vent length, date of emigration and survival for A. maculatum were all predicted by combinations of intra- and interspecific effects, but often showed negative relationships with A. annulatum metamorph traits and abundance. Size and date of metamorphosis were strongly correlated within each species, but in opposite patterns (negative for A. annulatum and positive for A. maculatum), suggesting that the two species use alternative strategies to enhance terrestrial survival and that these factors may influence their interactions. Our results match predictions from experimental studies that suggest recruitment is influenced by intra- and interspecific processes which are determined by phenological differences between species. Incorporating spatiotemporal variability when modeling population dynamics is necessary to understand the importance of phenology in species interactions, especially as shifts in phenology occur under climate change.
Sakata, Natsuko; Tamori, Yoichiro; Wakahara, Masami
Sex differentiation of gonads in amphibians is believed to be controlled genetically, but altered epigenetically or environmentally. When larvae of the salamander Hynobius retardatus were reared at defined temperatures from hatching to metamorphic stages, a high temperature (28 degrees C) induced exclusively female gonads (ovaries), whereas intermediate (20 and 23 degrees C) or lower (16 degrees C) temperatures produced a 1:1 sex ratio of the morphological gonads. The thermosensitive period was determined to be restricted from 15 to 30 days after hatching, just before or when sexual differentiation occurred. Hynobius P450 aromatase (P450arom) cDNA was isolated from adult gonads and the partial nucleotide or deduced amino acid sequences were determined, showing a high level of identity with various vertebrate species. The P450arom gene was expressed predominantly in the adult ovary and brain, weakly in testis, but not in other somatic organs. A typical sexual dimorphism in P450arom expression was detected in normally developing larvae by a quantitative competitive RT-PCR; strong expression in the female gonads but very weak in male gonads. The dimorphism was detected much earlier than the morphological sexual differentiation of the gonads. When larvae were reared at the female-producing temperature (28 degrees C), strong expression was detected in all the temperature-treated larvae, suggesting that P450arom was up-regulated, even in genetic males. Our results confirm the importance of the P450arom regulation in the sexual differentiation of gonads and demonstrate that an up-regulation of P450arom is involved in the process of temperature-sensitive sex reversal in this species.
Catherine D. McCusker
Full Text Available Regenerative medicine has reached the point where we are performing clinical trials with stem-cell-derived cell populations in an effort to treat numerous human pathologies. However, many of these efforts have been challenged by the inability of the engrafted populations to properly integrate into the host environment to make a functional biological unit. It is apparent that we must understand the basic biology of tissue integration in order to apply these principles to the development of regenerative therapies in humans. Studying tissue integration in model organisms, where the process of integration between the newly regenerated tissues and the ‘old’ existing structures can be observed and manipulated, can provide valuable insights. Embryonic and adult cells have a memory of their original position, and this positional information can modify surrounding tissues and drive the formation of new structures. In this Review, we discuss the positional interactions that control the ability of grafted cells to integrate into existing tissues during the process of salamander limb regeneration, and discuss how these insights could explain the integration defects observed in current cell-based regenerative therapies. Additionally, we describe potential molecular tools that can be used to manipulate the positional information in grafted cell populations, and to promote the communication of positional cues in the host environment to facilitate the integration of engrafted cells. Lastly, we explain how studying positional information in current cell-based therapies and in regenerating limbs could provide key insights to improve the integration of cell-based regenerative therapies in the future.
Kumar, Anoop; Nevill, Graham; Brockes, Jeremy P; Forge, Andrew
Limb regeneration in salamanders proceeds by formation of the blastema, a mound of proliferating mesenchymal cells surrounded by a wound epithelium. Regeneration by the blastema depends on the presence of regenerating nerves and in earlier work it was shown that axons upregulate the expression of newt anterior gradient (nAG) protein first in Schwann cells of the nerve sheath and second in dermal glands underlying the wound epidermis. The expression of nAG protein after plasmid electroporation was shown to rescue a denervated newt blastema and allow regeneration to the digit stage. We have examined the dermal glands by scanning and transmission electron microscopy combined with immunogold labelling of the nAG protein. It is expressed in secretory granules of ductless glands, which apparently discharge by a holocrine mechanism. No external ducts were observed in the wound epithelium of the newt and axolotl. The larval skin of the axolotl has dermal glands but these are absent under the wound epithelium. The nerve sheath was stained post-amputation in innervated but not denervated blastemas with an antibody to axolotl anterior gradient protein. This antibody reacted with axolotl Leydig cells in the wound epithelium and normal epidermis. Staining was markedly decreased in the wound epithelium after denervation but not in the epidermis. Therefore, in both newt and axolotl the regenerating axons induce nAG protein in the nerve sheath and subsequently the protein is expressed by gland cells, under (newt) or within (axolotl) the wound epithelium, which discharge by a holocrine mechanism. These findings serve to unify the nerve dependence of limb regeneration.
Many workers interested in the mechanics and kinematics of undulatory aquatic locomotion have examined swimming in fishes that use a carangiform or subcarangiform mode. Few empirical data exist describing and quantifying the movements of elongate animals using an anguilliform mode of swimming. Using high-speed video, I examine the axial undulatory kinematics of an elongate salamander, Siren intermedia, in order to provide data on how patterns of movement during swimming vary with body position and swimming speed. In addition, swimming kinematics are compared with those of other elongate vertebrates to assess the similarity of undulatory movements within the anguilliform locomotor mode. In Siren, most kinematic patterns vary with longitudinal position. Tailbeat period and frequency, stride length, Froude efficiency and the lateral velocity and angle of attack of tail segments all vary significantly with swimming speed. Although swimming speed does not show a statistically significant effect on kinematic variables such as maximum undulatory amplitude (which increases non-linearly along the body), intervertebral flexion and path angle, examination of the data suggests that speed probably has subtle and site-specific effects on these variables which are not detected here owing to the small sample size. Maximum lateral displacement and flexion do not coincide in time within a given tailbeat cycle. Furthermore, the maximum orientation (angle with respect to the animal's direction of forward movement) and lateral velocity of tail segments also do not coincide in time. Comparison of undulatory movements among diverse anguilliform swimmers suggests substantial variation across taxa in parameters such as tailbeat amplitude and in the relationship between tailbeat frequency and swimming speed. This variation is probably due, in part, to external morphological differences in the shape of the trunk and tail among these taxa.
Full Text Available Computer simulation has been used to investigate several aspectsof locomotion in salamanders. Here we introduce a three-dimensionalforward dynamics mechanical model of a salamander, with physicallyrealistic weight and size parameters. Movements of the four limbs and ofthe trunk and tail are generated by sets of linearly modeled skeletalmuscles. In this study, activation of these muscles were driven byprescribed neural output patterns. The model was successfully used tomimic locomotion on level ground and in water. We compare thewalking gait where a wave of activity in the axial muscles travelsbetween the girdles, with the trotting gait in simulations usingthe musculo-mechanical model. In a separate experiment, the model is usedto compare different strategies for turning while stepping; either bybending the trunk or by using side-stepping in the front legs. We foundthat for turning, the use of side-stepping alone or in combination withtrunk bending, was more effective than the use of trunk bending alone. Weconclude that the musculo-mechanical model described here together with aproper neural controller is useful for neuro-physiological experiments insilico.
Full Text Available The discovery and subsequent description of cryptic biodiversity is often challenging, especially for groups that have undergone rapid lineage accumulation in the relatively recent past. Even without formal descriptions, understanding genetic diversity patterns as they relate to underlying ecological or historical processes can be important for conservation. The dusky salamanders of the genus Desmognathus, with 20 described species, comprise the second largest genus of plethodontid salamanders in the eastern United States. However, due to the presence of high genetic diversity and relatively few morphological synapomorphies, the number of species is likely to increase. For the three nominal species within the D. quadramaculatus species complex, including D. quadramaculatus, D. folkertsi, and D. marmoratus, we used a portion of the mitochondrial genome and nuclear markers in the form of amplified fragment length polymorphisms (AFLP to uncover spatial patterns of genetic diversity. Within D. quadramaculatus and D. marmoratus, we uncovered four well-supported lineages with the mitochondrial sequences; phylogeographic patterns were not congruent with the AFLP data. Both sets of markers identified a clear isolation by stream distance. Using multiple regressions, we found that historical river drainages and terrestrial ecoregions explained the phylogeographic patterning we observed for D. quadramaculatus.
Julio A. Lemos-Espinal
Full Text Available Several species of salamander in the genus Ambystoma occur in the mountains surrounding Mexico City and are considered at risk of extinction. However, little is known about their ecology and natural history. The Toluca Stream Siredon (Ambystoma rivulare is classified as “Data Deficient” by the IUCN, and considered “Threatened” under Mexican law. From October 2013 to September 2014, we examined the diet of larval A. rivulare from a stream on the Volcán Nevado de Toluca in Mexico to provide insight into the suitability of the habitat to support this population of salamanders. Ostracods accounted for approximately 90% of all prey items consumed by larval A. rivulare. The number of ostracods found in stomachs increased with individual body size, but the proportion of ostracods in stomachs did not vary with body size. Nematodes were observed in approximately one third of the stomachs we examined. The diversity of prey in the diet of A. rivulare in the stream we studied is low and dominated by a single prey taxon, ostracods. Our results suggest that if environmental conditions in the stream change such that ostracods are negatively affected then the long-term persistence of this population of A. rivulare might be in jeopardy.
Baxter, Leilan; Brain, Richard A; Hosmer, Alan J; Nema, Mohini; Müller, Kirsten M; Solomon, Keith R; Hanson, Mark L
Embryonic growth of the yellow-spotted salamander (Ambystoma maculatum) is enhanced by the presence of the green alga Oophila amblystomatis, in the egg capsule. To further assess potential impacts of herbicides on this relationship, A. maculatum egg masses were exposed to atrazine (0-338 μg/L) until hatching (up to 66 days). Exposure to atrazine reduced PSII yield of the symbiotic algae in a concentration-dependent manner, but did not significantly affect visible algal growth or any metrics associated with salamander development. Algal cells were also cultured in the laboratory for toxicity testing. In the 96-h growth inhibition test (0-680 μg/L), ECx values were generally greater than those reported for standard algal test species. Complete recovery of growth rates occurred within 96-h of transferring cells to untreated media. Overall, development of A. maculatum embryos was not affected by exposure to atrazine at concentrations and durations exceeding those found in the environment.
Blassberg, Robert A; Garza-Garcia, Acely; Janmohamed, Azara; Gates, Phillip B; Brockes, Jeremy P
The GPI-anchor is an established determinant of molecular localisation and various functional roles have been attributed to it. The newt GPI-anchored three-finger protein (TFP) Prod1 is an important regulator of cell behaviour during limb regeneration, but it is unclear how it signals to the interior of the cell. Prod1 was expressed by transfection in cultured newt limb cells and activated transcription and expression of matrix metalloproteinase 9 (MMP9) by a pathway involving ligand-independent activation of epidermal growth factor receptor (EGFR) signalling and phosphorylation of extracellular regulated kinase 1 and 2 (ERK1/2). This was dependent on the presence of the GPI-anchor and critical residues in the α-helical region of the protein. Interestingly, Prod1 in the axolotl, a salamander species that also regenerates its limbs, was shown to activate ERK1/2 signalling and MMP9 transcription despite being anchorless, and both newt and axolotl Prod1 co-immunoprecipitated with the newt EGFR after transfection. The substitution of the axolotl helical region activated a secreted, anchorless version of the newt molecule. The activity of the newt molecule cannot therefore depend on a unique property conferred by the anchor. Prod1 is a salamander-specific TFP and its interaction with the phylogenetically conserved EGFR has implications for our view of regeneration as an evolutionary variable.
Heredia-Bobadilla, Rosa-Laura; Monroy-Vilchis, Octavio; Zarco-González, Martha M; Martínez-Gómez, Daniel; Mendoza-Martínez, Germán David; Sunny, Armando
Human activities are affecting the distribution of species worldwide by causing fragmentation and isolation of populations. Isolation and fragmentation lead to populations with lower genetic variability and an increased chance of inbreeding and genetic drift, which results in a loss of biological fitness over time. Studies of the genetic structure of small and isolated populations are critically important for management and conservation decisions. Ambystoma rivulare is a micro-endemic Mexican mole salamander from central Mexico. It is found in the most ecologically disturbed region in Mexico, the Trans-Mexican Volcanic Belt. The goal of this study of the population genetics of the micro-endemic mole salamander was to provide information to be used as a basis for future research and conservation planning of this species and other species of the Ambystoma genus in Mexico. The structural analysis found two subpopulations, one for each river sampled, with no signs of admixture and very high levels of genetic differentiation. Medium to high levels of heterozygosity and few alleles and genotypes were observed. Evidence of an ancestral genetic bottleneck, low values of effective population size, small inbreeding coefficients, and low gene flow were also found.
Pietsch, P; Schneider, C W
A wide variety of visual functions show increases attributable to binocularity, and the question pursued here was whether a second eye enhances the visually stimulated skin blanching reaction of the larval salamander. Dermal melanin spots (produced by the aggregations of melanosomes within dermal melanophores and which contract or expand to lighten or darken the skin) were measured in eyeless (controls), one-eyed and two-eyed Ambystoma punctatum larvae after chronic adaptation of the subjects to a white background (i.e., stimulus conditions for maximum blanching). The eyeless subjects showed no blanching (thus remained dark) in white cups, and they exhibited melanin spots 7 or 8 times the size of those of the other two groups. All one-eyed or two-eyed subjects exhibited blanching reactions; planometric comparison revealed a significantly larger melanin spot area for one-eyed than for two-eyed animals; i.e., the binocular condition permitted greater contraction of the pigment spots than did the monocular condition. Analytical data compared favorably with independently ascertained pigmentation indices. The results indicate that a second eye quantitatively elevates the blanching maximum of a larval salamander.
Harischandra, Nalin; Cabelguen, Jean-Marie; Ekeberg, Orjan
Computer simulation has been used to investigate several aspects of locomotion in salamanders. Here we introduce a three-dimensional forward dynamics mechanical model of a salamander, with physically realistic weight and size parameters. Movements of the four limbs and of the trunk and tail are generated by sets of linearly modeled skeletal muscles. In this study, activation of these muscles were driven by prescribed neural output patterns. The model was successfully used to mimic locomotion on level ground and in water. We compare the walking gait where a wave of activity in the axial muscles travels between the girdles, with the trotting gait in simulations using the musculo-mechanical model. In a separate experiment, the model is used to compare different strategies for turning while stepping; either by bending the trunk or by using side-stepping in the front legs. We found that for turning, the use of side-stepping alone or in combination with trunk bending, was more effective than the use of trunk bending alone. We conclude that the musculo-mechanical model described here together with a proper neural controller is useful for neuro-physiological experiments in silico.
Lamb, T D; Matthews, H R; Torre, V
The suction-electrode technique was used to monitor the photocurrent of isolated retinal rods from the tiger salamander, by drawing in the light-sensitive outer segment, or sometimes the inner segment. Calcium buffers or other agents were then introduced into the rod cytoplasm by the 'whole-cell patch-clamp' technique. A patch pipette was sealed against the region of the rod protruding from the suction pipette (usually the inner segment), and the membrane patch was ruptured to obtain a whole-cell recording. Several lines of evidence indicated that the pipette contents diffused into the outer segment, and showed that the cell could be adequately voltage clamped. With only trace quantities of chelator in the patch pipette (to bind stray calcium), a gradual decline of the dark current and slowing of responses was usually observed over a period of 10-20 min after rupture of the patch. When the patch pipette contained no added calcium and 10 mM of the calcium chelator BAPTA (1,2-bis(o-aminophenoxy)ethane-N,N,N',N'-tetraacetic acid) (free Ca2+ ca. 10(-9) M) rupture of the patch led, over a period of a few minutes, to an increase in mean dark current, an increased duration of responses, a substantial increase in flash sensitivity, and a pronounced overshoot in the recovery phase, but with virtually no change in the rising phase of the response to bright flashes. Similar results were obtained when EGTA was used in place of BAPTA, and also in the few cases when successful rupture of the outer segment membrane was obtained. With the free calcium concentration in the patch pipette buffered to the higher level of 1 microM (with 10 mM-Ca2+/11 mM-BAPTA) the results were qualitatively similar to those obtained with BAPTA alone, except that the mean dark current did not increase. This is consistent with a resting free calcium concentration in darkness in the region of 1 microM. In the presence of bright steady illumination with BAPTA in the cell the suppression of outer segment
Full Text Available L’article examine l’idée d’événement à travers la figure de la salamandre dans l’œuvre d’Yves Bonnefoy. L’image, saisissante en elle-même, est rendue plus significative encore par le fait qu’elle franchit les frontières séparant habituellement les genres littéraires. Son apparition à la fois dans un essai et dans un cycle de poèmes tend à l’ancrer dans la réalité d’un événement vécu. Nous proposons de voir dans la juxtaposition des deux textes non seulement une narration qui prend l’événement vécu en charge, mais l’agencement même des possibilités permettant de reconnaître l’événement comme tel.This paper addresses the notion of event through the prism of Yves Bonnefoy’s salamander trope. This arresting image is all the more memorable because it crosses the boundaries of generic separations, appearing both in an essay as well as in an early poetry collection. The fact that an identical image should be reiterated in such formally disparate structures seems to point insistently to its reality as a material event. I argue that juxtaposing the two texts reveals both a narration recounting an event and, simultaneously, the ordering of possibilities leading to its recognition as event.El artículo examina la idea del acontecimiento a través de la figura de la salamandra en la obra de Yves Bonnefoy. La imagen, sorprendente en sí misma, se vuelve más significativa aún debido al hecho de que cruza las fronteras que separan generalmente los géneros literarios. Su presencia tanto en un ensayo como en un ciclo de poemas tiende a situarla en la realidad de un acontecimiento vivido. En la yuxtaposición de los dos textos proponemos ver no sólo una narración que asume el acontecimiento vivido, sino también la disposición misma de las posibilidades que permiten reconocer al acontecimiento como tal.
王苗苗; 罗庆华; 王海磊; 宋英杰; 王建文; 向建国
The oil was extracted from the tail adipose tissue of Giant Salamander by enzymatic hydrolysis method so as to seek a new way for the utilization of Chinese Giant Salamander resource. Then the physi-cochemical properties and fatty acid composition of Giant Salamander tail oil were analyzed. The results showed that the optimal enzymatic extraction conditions of Giant Salamander tail oil were obtained as fol-lows:dosage of enzyme 0. 3%,ratio of solid to liquid 1∶1. 0, enzymolysis time 1. 0 h,enzymolysis temper-ature 60℃,pH 6. 5. Under the optimal conditions, the extraction rate was 98. 53%. The Giant Salaman-der tail oil extracted reached the first grade standard of crude fish oil ( SC/T 3502—2000 ) . 17 kinds of fatty acids were detected from the Giant Salamander tail oil,and the contents of MUFA and PUFA were 49. 42% and 29. 35% respectively.%为寻求我国大鲵资源利用新途径,并建立大鲵尾部油的最佳提取方法及工艺,采用酶解法从大鲵尾部脂肪组织中提取大鲵尾部油,并对所得的大鲵尾部油进行理化性质及脂肪酸组成分析。结果表明,酶解法提取大鲵尾部油的最佳工艺条件为：酶添加量0.3%,料液比1∶1.0,酶解时间1.0 h,酶解温度60℃,pH 6.5；在最佳条件下,提取率为98.53%。所提的大鲵尾部油达到我国水产行业SC/T 3502—2000粗鱼油的一级标准。酶解法所提大鲵尾部油中检测出17种脂肪酸,单不饱和脂肪酸(MUFA)含量为49.42%,多不饱和脂肪酸(PUFA)含量为29.35%。
Mohammad M. Al-Qattan
Full Text Available nAG (newt-Anterrior Gradient protein is the key mediator of regrowth of amputated limbs in salamanders. In a previous work in our lab, a new nAG gene (suitable for humans was designed and cloned. The cloned vector was transfected into primary human fibroblasts. The expression of nAG in human primary fibroblasts was