Parasitism of Hylodes phyllodes (Anura: Cycloramphidae) by Hannemania sp (Acari: Trombicuudae) in an area of Atlantic Forest, Ilha Grande, southeastern Brazil

We studied some parameters of the parasitism by the mite Hannemonia sp. on the endemic frog Hylodes phyllodes in the Atlantic Forest of Ilha Grande (Rio de Janeiro State, Brazil). Prevalence, mean abundance, mean intensity and total intensity of infestation, body regions infected, and host sexual differences in parasitism rate of the larvae of Hannemania sp. on individuals of H. phyllodes were assessed. Prevalence was 86.5% (N = 37; total of 1,745 larvae of Hannemania sp) for male hosts and 91.7% (N = 12; total of 330 larvae) for female hosts, with no significant difference between the sexes. Overall prevalence of Hannemania sp. on H. phyllodes was 87.7%. Mean intensity of infestation in males (54.5 +/- 42.5; range 1-173 larvae) was higher than in females (29.9 +/- 47.6; range 1-166). We conclude that the rates of intensity, abundance, and prevalence of Hannemania larvae parasitizing Hylodes phyllodes at Ilha Grande were considerably high, suggesting that this species of anuran constitutes a relevant host for this mite species to complete its life-cycle in the area. Differences between males and females in infestation rates probably reflect their differential use of space in the forest.     

Systematics of dusky salamanders, Desmognathus (Caudata: Plethodontidae), in the mountain and Piedmont regions of Virginia and North Carolina, USA

We analysed mitochondrial (cytochrome  b ) nucleotide sequences, nuclear allozyme markers, and morphometric characters to investigate species boundaries and phylogenetic relationships among dusky salamanders ( Desmognathus ) in the southern Blue Ridge and adjacent Piedmont Physiographic Provinces of Virginia and North Carolina. Our results revealed four distinct mitochondrial DNA clades that are also characterized by distinct allozyme markers. One clade consists of sequences derived from populations distributed from New England to south-western Virginia that are referable to Desmognathus fuscus Rafinesque, 1820, although there is considerable sequence and allozyme divergence within this clade. A second clade consists of sequences derived from populations referable to Desmognathus planiceps Newman, 1955, a form that we resurrect from its long synonymy under D. fuscus . Desmognathus planiceps and D. fuscus also differ in mandibular tooth shape. Two other cytochrome  b sequences recovered from populations along the Blue Ridge escarpment in southern Virginia are quite divergent from those of the previous two clades, and these populations may represent yet another distinct species. Sequences from a population in the Brushy Mountains in the Piedmont of northern North Carolina are similar to those of Desmognathus carolinensis . Population groupings indicated by allozyme data generally correspond to the cytochrome  b clades. Cryptic diversity in Appalachian desmognathan salamanders clearly requires further study. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 115–130.     

Evaluating the effects of urbanisation on salamander abundances using a before‐after control‐impact design

1. Urbanisation represents a significant threat to semi‐aquatic amphibian populations, especially stream‐dwelling salamanders. Although studies of urbanisation effects on amphibians have been conducted, there is an urgent need to follow populations over longer time periods, account for imperfect detection and determine the response time to urbanisation. Consequently, we used a before‐after control‐impact (BACI) study design to estimate changes in abundances of larval and adult salamanders in streams affected by urbanisation. 2. From 2005 to 2009, we used standard sampling techniques to obtain a count of salamanders in 13 first‐order streams that underwent urbanisation of their catchments after the first year of sampling. Simultaneously, we counted salamanders in 17 streams that experienced no disturbance within stream catchments. Additionally, we measured environmental variables at each stream. 3. We used Royle’s binomial mixture model to estimate annual mean abundances and individual detection probabilities, and Bayesian inference was used to estimate population parameters for each stage and species. 4. Although mean abundance estimates varied among years in control and urbanised streams, we found that urbanisation had a negative effect on larval and adult salamander abundances. Larval salamander abundances at sites 1 year after urbanisation were significantly lower than abundances from control sites. Abundances of adult two‐lined salamanders (Eurycea cirrigera) at urbanised sites were lower than abundances at control sites 2 years post‐urbanisation, and adult dusky salamander (Desmognathus fuscus) abundances at urbanised sites were lower than abundances at control sites 3 years post‐urbanisation. Maximum conductivity, sedimentation level and maximum stream channel width differed between urban and non‐urban streams. 5. Our results suggest that stream‐dwelling salamanders exhibit little resistance to urbanisation. Our study also highlights the use of the BACI design to study how urbanisation affects populations in semi‐aquatic habitats. We emphasise that inferences regarding urbanisation effects on population response may be compromised unless urban populations are compared to populations in control sites, especially for species in which populations fluctuate.     

Helminth and leech community structure in tadpoles and caudatan larvae of two amphibian species from Western Nebraska

Currently no comparative studies exist on helminth and leech community structure among sympatric anuran tadpoles and salamander larvae. During June-August 2007-2009, we examined 50 bullfrog tadpoles, Rana catesbeiana , 50 barred tiger salamander larvae, Ambystoma mavortium , and 3 species of snails from Nevens Pond, Keith County, Nebraska for helminth and leech infections. The helminth and leech compound community of this larval amphibian assemblage consisted of at least 7 species, 4 in bullfrog tadpoles and 4 in barred tiger salamander larvae. Bullfrog tadpoles were infected with 2 species of nematodes ( Gyrinicola batrachiensis and Spiroxys sp.) and 2 types of metacercariae ( Telorchis sp. and echinostomatids), whereas barred tiger salamander larva were infected with 1 species of leech ( Placobdella picta ), 2 species of adult trematodes ( Telorchis corti and Halipegus sp.), and 1 species of an unidentified metacercaria. The component community of bullfrog tadpoles was dominated by helminths acquired through active penetration, or incidentally ingested through respiratory currents, or both, whereas the component community of larval salamanders was dominated by helminths acquired through ingestion of intermediate hosts (χ2 = 3,455.00, P < 0.00001). Differences in amphibian larval developmental time (2-3 yr for bullfrog tadpoles versus 2-5 mo for salamander larvae), the ephemeral nature of intermediate hosts in Nevens Pond, and the ability of bullfrog tadpole to eliminate echinostome infections had significant effects on mean helminth species richness among amphibian species and years (t = 12.31, P < 0.0001; t = 2.09, P = 0.04). Differences in herbivorous and carnivorous diet and time to metamorphosis among bullfrog tadpoles and barred tiger salamander larvae were important factors in structuring helminth communities among the larval stages of these 2 sympatric amphibian species, whereas size was important in structuring helminth and leech communities in larval salamanders, but not in bullfrog tadpoles.     

The courtship behaviour of the spotted dusky salamander, Desmognathus fuscus conanti (Amphibia: Caudata: Plethodontidae)

The courtship behaviour of the North American southern dusky salamander, Desmognathus fuscus conanti (Plethodontidae), is described for the first time. The male is the most active participant, as is typical for caudate amphibians, and much of his behaviour can be understood in terms of orientation to and stimulation (or persuasion) of the female. The latter is largely accomplished by the transfer of courtship pheromones from the male's mental (chin) gland. The orientation and persuasion phases of courtship appear necessary if sperm transfer by means of a spermatophore deposited upon the substratum is to be successful. The courtship of D. fuscus conanti is qualitatively identical to that of D. santeetlah , with which it is sexually compatible in the laboratory and with which it may hybridize in nature. However, a broader comparison of species in the genus Desmognathus reveals that the greatest interspecific differences occur during the orientation and persuasion phases of courtship. A complete understanding of the evolution of courtship behaviour in desmognathine salamanders awaits availability of ethograms for all other taxa and a robust, independent phytogeny for the entire subfamily.     

Divergence of mate recognition systems among conspecific populations of the plethodontid salamander Desmognathus santeetlah (Amphibia: Plethodontidae)

The disjunct geographic distribution of plethodontid salamanders of the genus Desmognathus provides excellent opportunities to test whether populations diverge in mate recognition systems so as to result in sexual incompatibility. Previous work has shown that sexual incompatibility may exist both among conspecific populations and between different species in this genus. Populations of the Santeetlah dusky salamander, Desmognathus santeetlah , are found at higher elevations in three mountain ranges in the southern Appalachian Mountains of eastern North America, where they are sympatric with congeneric species of similar body size. We report that statistically significant levels of sexual incompatibility exist among populations of D. santeetlah collected from across the species' range. Contrary to popular belief, but in accord with a growing body of empirical literature, aspects of mate recognition systems can diverge among conspecific populations of salamanders, resulting in incidental sexual incompatibility.     

Responses of a plethodontid salamander to conspecific and congeneric intruders

After inhabiting their home sites for a few days, both males and females of the salamander Desmognathus monticola directed agonistic behaviour toward intruding salamanders. Resident D. monticola were significantly more aggressive toward D. fuscus intruders than toward conspecific ones, although both conspecific and congeneric intruders were frequently attacked. Responses of D. monticola residents were independent of sex, and no difference was found in the behaviour of residents toward conspecific male and female intruders. Small D. fuscus intruders were attacked and injured more frequently than similar-sized D. monticola intruders and, in general, smaller-sized salamanders were the victims of aggressive attacks more frequently than larger ones. The proportions of bites on the head and tail of intruders were similar, although injuries were much more frequent to the tail than to the head. This species-related and size-related agonistic behaviour may be functional in maintaining the spatial distribution of species in communities of Desmognathus salamanders.     

DO LARVAL TRAITS RE‐EVOLVE? EVIDENCE FROM THE EMBRYOGENESIS OF A DIRECT‐DEVELOPING SALAMANDER,PLETHODON CINEREUS

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.     

Diet composition of two larval headwater stream salamanders and spatial distribution of prey

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Salamander diversity alters stream macroinvertebrate community structure

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Cytophotometric evidence of variation in genome size of desmognathine salamanders

The amount of DNA per haploid genome, the C-value, is often directly correlated with nuclear and cell volume, but inversely correlated with cell replication rate. Also, rates of cellular growth sometimes appear to be correlated with organismal developmental rates and life history patterns. Among vertebrates, salamanders exhibit the greatest variation in genome size. In the present study we have examined interspecific and intraspecific variation in blood cell DNA levels in the genus Desmognathus, which shows greater variation in life history traits than any other salamander genus. Specimens of Desmognathus quadramaculatus, D. monticola, D. ochrophaeus and D. wrighti were collected from nature at two localities in the southern Appalachian Mountains. Estimates of genome size in pg of DNA were obtained from blood smears by DNA-Feulgen cytophotometry, using erythrocyte nuclei of Xenopus laevis as an internal reference standard of 6.35 pg DNA per cell. C-values of Desmognathus are the smallest in the order Caudata. Although significant variation in DNA levels was found among the four species, the differences were small, and do not support previously proposed relationships between C-value and life-history variation.     

Helminth community structure of sympatric eastern American toad, Bufo americanus americanus, northern leopard frog, Rana pipiens, and blue-spotted salamander, Ambystoma laterale, from southeastern Wisconsin

One-hundred twelve amphibians, including 51 blue-spotted salamanders, Ambystoma laterale, 30 eastern American toads, Bufo americanus americanus, and 31 northern leopard frogs, Rana pipiens, were collected during April-October 1996 from Waukesha County, Wisconsin and examined for helminth parasites. The helminth compound community of this amphibian assemblage consisted of at least 10 species: 9 in American toads, 8 in leopard frogs, and 3 in blue-spotted salamanders. American toads shared 7 species with leopard frogs, and 2 species occurred in all 3 host species. Although there was a high degree of helminth species overlap among these sympatric amphibians, statistically significant differences were found among host species and percent of indirect or direct-life cycle parasites of amphibian species individual component communities (chi2 = 1,015, P < 0.001). American toads had a higher relative abundance of nematodes, 59%, than larval cestodes, 31%, and larval and adult trematodes, 10%, whereas leopard frogs had a higher relative abundance of larval cestodes, 71.3%, and larval and adult trematodes, 25.3%, than nematodes 3.4%. This is related to ecological differences in habitat and dietary preferences between these 2 anuran species. Helminth communities of blue-spotted salamanders were depauperate and were dominated by larval trematodes, 94%, and few nematodes, 6%. Low helminth species richness in this host species is related to this salamander's relatively small host body size, smaller gape size, lower vagility, and more fossorial habitat preference than the other 2 anuran species. Adult leopard frogs and toads had significantly higher mean helminth species richness than metamorphs, but there was no significant difference in mean helminth species richness among adult and metamorph blue-spotted salamanders. Considering adult helminths, the low species richness and low vagility of caudatans as compared with anurans suggest that local factors may be more important in structuring caudatan helminth communities of salamanders than of anuran hosts. Helminth species infecting salamanders may be more clumped in their geographic distribution as compared with anurans, and the role of other hosts and their parasites at the compound community level may be important in structuring helminth communities of salamanders.     

A New Species of Eimeria (Apicomplexa: Eimeriidae) from the marbled salamander, Ambystoma opacum (Caudata: Ambystomatidae), from northern Louisiana

Between December 2002 and June 2004, 10 marbled salamanders, Ambystoma opacum, were examined for coccidian parasites. Salamanders were collected in Bradley (n = 2), Little River (n = 1), Miller (n = 1), and Sevier (n = 1) Counties, Arkansas; Webster Parish, Louisiana (n = 2); and Bowie (n = 1) and Nacogdoches (n = 2) Counties, Texas. Two of 10 (20%) A. opacum from Louisiana harbored an undescribed species of Eimeria. Oocysts of Eimeria trauthi n. sp. were ellipsoidal, 36.6 x 33.1 (33-40 x 29-37) microm, with a thin, single-layered wall; shape index 1.1. Polar granule(s) and micropyle were absent. Oocyst residuum was composed of hundreds of loosely packed homogenous granules of various sizes enclosing a vacuole. Sporocysts were elongate-ellipsoidal, 20.8 x 8.1 (19-22 x 7-9) microm; shape index 2.6. Sporocyst residuum was spherical and composed of a cluster of granules often membrane-bound. This is the first time a coccidium has been reported from an amphibian species in Louisiana and the second time a coccidium has been described from this salamander host. In addition, the following 26 salamanders from various counties in Arkansas, Oklahoma, and Texas were surveyed during the study period and were negative for coccidia: Ambystomatidae, 4 spotted salamanders (Ambystoma maculatum) and 7 mole salamanders (Ambystoma talpoideum); Cryptobranchidae, 4 Ozark hellbenders (Cryptobranchus alleganiensis bishopi); Plethodontidae, 6 spotted dusky salamanders (Desmognathus conanti) and 3 many-ribbed salamanders (Eurycea multiplicata multiplicata); and Salamandridae, 2 central newts (Notophthalmus viridescens louisianensis).     

Frequency of interspecific mating in salamanders of the plethodontid genus Desmognathus: different experimental designs may yield different results

Heterosexual courtship trials were staged between dusky salamanders of two species ( Desmognathus fuscus and D. santeetlah ) from two allopatric populations in the southern Appalachian Mountains of eastern North America. Two experiments were conducted in order to test for effects of experimental design on estimates of the strength of sexual isolation between these species. In Experiment 1, frequencies of interspecific mating were determined when: (1) a male was presented with a conspecific and a heterotypic female successively; and (2) both types of females were presented simultaneously to a male. The frequency of interspecific mating by male D. fuscus was greatest when females were presented successively. The frequency of interspecific mating by male D. santeetlah was not influenced by method of female presentation. An index of sexual isolation of +0.29 was obtained between D. fuscus and D. santeetlah when females were presented successively, and of +0.63 when females were presented simultaneously. In Experiment 2, interspecific mating frequencies were determined in courtship arenas of two different sizes. No significant effect of arena size on frequency of interspecific mating was found for D. fuscus. In D. santeetlah , a trend towards greater mating frequency in small arenas was almost significant. These results suggest that interspecific mating frequencies (and thus estimates of the strength of sexual isolation between populations or species) may be sensitive to the experimental design employed. The validity and generality of this conclusion for other animals is an urgent, empirical issue.     

Feather mites (Acari: Astigmata) and body condition of their avian hosts: a large correlative study

Feather mites are arthropods that live on or in the feathers of birds, and are among the commonest avian ectosymbionts. However, the nature of the ecological interaction between feather mites and birds remains unclear, some studies reporting negative effects of feather mites on their hosts and others reporting positive or no effects. Here we use a large dataset comprising 20 189 measurements taken from 83 species of birds collected during 22 yr in 151 localities from seven countries in Europe and North Africa to explore the correlation between feather mite abundance and body condition of their hosts. We predicted that, if wing‐dwelling feather mites are parasites, a negative correlation with host body condition should be found, while a mutualistic interaction should yield positive correlation. Although negative relationships between feather mite abundance and host body condition were found in a few species of birds, the sign of the correlation was positive in most bird species (69%). The overall effect size was only slightly positive (r =0.066). The effect of feather mite abundance explained <10% of variance in body condition in most species (87%). Results suggest that feather mites are not parasites of birds, but rather that they hold a commensalistic relationship where feather mites may benefit from feeding on uropygial gland secretions of their hosts and birds do not seem to obtain a great benefit from the presence of feather mites.     

Leaf domatia and foliar mite abundance in broadleaf deciduous forest of north Asia

Plant morphology may be shaped, in part, by the third trophic level. Leaf domatia, minute enclosures usually in vein axils on the leaf underside, may provide the basis for protective mutualism between plants and mites. Domatia are particularly frequent among species of trees, shrubs, and vines in the temperate broadleaf deciduous forests in north Asia where they may be important in determining the distribution and abundance of mites in the forest canopy. In lowland and montane broadleaf deciduous forests at Kwangn;akung and Chumbongsan in Korea, we found that approximately half of all woody species in all forest strata, including many dominant trees, have leaf domatia. Pooling across 24 plant species at the two sites, mites occupied a mode of 60% (range 20-100%) of domatia and used them for shelter, egg-laying, and development. On average, 70% of all active mites and 85% of mite eggs on leaves were found in domatia; over three-quarters of these were potentially beneficial to their hosts. Further, mite abundance and reproduction (expressed as the proportion of mites at the egg stage) were significantly greater on leaves of species with domatia than those without domatia in both forests. Effects of domatia on mite abundance were significant only for predaceous and fungivorous mite taxa; herbivore numbers did not differ significantly between leaves of species with and without domatia. Comparable patterns in broadleaf deciduous forest in North America and other biogeographic regions suggest that the effect of leaf domatia on foliar mite abundance is general. These results are consistent with several predictions of mutualism between plants and mites, and indicate that protective mutualisms may be frequent in the temperate zone.     

Larval parasitism of spring-dwelling alpine water mites (Hydrachnidia,Acari): a study with particular reference to chironomid hosts

During faunistic investigations on spring habitats in the alpine National Park Berchtesgaden (Bavaria, Germany), water mites were found to be the group with the highest share of species strongly adapted to springs. At four sample sites at two spring complexes, insect emergence was screened for parasitism by larval water mites. A total of at least 36 host species were recorded as being parasitized by 19 water mite species. As in many other habitats, the most important host taxon was shown to be the nematoceran family Chironomidae, both in the number of species and individuals parasitized. Likewise, the number of water mite species attached to chironomids was high. Further host species were found among the Plecoptera, Trichoptera, Coleoptera, Limoniidae and Empididae (Diptera). These taxa were only parasitized by a single water mite species in each case. For 13 mite species, new hosts were recorded for the first time. For another six species, the known host spectrum could be confirmed and/or supplemented. The parasitological data presented (e.g., prevalences, selected attachment sites on the host, larval phenology, intensity of parasitism) provide, in most cases, basic information concerning previously unknown parasite–host associations. At this time, the reason for the strong crenobiosis in water mites cannot be explained by their parasitism.     

Dusky salamanders (Desmognathus, Plethodontidae) from the Coastal Plain: multiple independent lineages and their bearing on the molecular phylogeny of the genus

Recent phylogenetic reassessment of the lungless salamanders (Plethodontidae) confirmed a major life-history reversal-from direct development to an aquatic larval stage-in the dusky salamanders (Desmognathus) of eastern North America. This reversal initiated high rates of lineage accumulation, reputedly generating the species richness and ecological breath that now characterize Desmognathus. Certain important aspects of the radiation, e.g., ecomorphological evolution, have been identified through intense sampling effort of Appalachian Highland lineages. However, the research preoccupation on montane species has left overlooked a significant component of dusky salamander distribution-the Coastal Plain. We present the first molecular phylogeny for Desmognathus to incorporate extensive coverage from the Atlantic and Gulf coastal plains. We examined 38 Coastal Plain populations in conjunction with 45 additional populations, representing 16 of the 19 nominal species. Bayesian analysis of 88 mitochondrial cox1 haplotypes diagnosed eight independent population lineages within the Coastal Plain, a number at odds with the region's three currently recognized species. Desmognathus has apparently experienced a complex biogeographic history in this physiographic region, one involving multiple invasions and several ecological transitions from lotic to lentic habitats.     

Feather mite abundance varies but symbiotic nature of mite‐host relationship does not differ between two ecologically dissimilar warblers

Feather mites are obligatory ectosymbionts of birds that primarily feed on the oily secretions from the uropygial gland. Feather mite abundance varies within and among host species and has various effects on host condition and fitness, but there is little consensus on factors that drive variation of this symbiotic system. We tested hypotheses regarding how within‐species and among‐species traits explain variation in both (1) mite abundance and (2) relationships between mite abundance and host body condition and components of host fitness (reproductive performance and apparent annual survival). We focused on two closely related (Parulidae), but ecologically distinct, species: Setophaga cerulea (Cerulean Warbler), a canopy dwelling open‐cup nester, and Protonotaria citrea (Prothonotary Warbler), an understory dwelling, cavity nester. We predicted that feather mites would be more abundant on and have a more parasitic relationship with P. citrea, and within P. citrea, females and older individuals would harbor greater mite abundances. We captured, took body measurements, quantified feather mite abundance on individuals’ primaries and rectrices, and monitored individuals and their nests to estimate fitness. Feather mite abundance differed by species, but in the opposite direction of our prediction. There was no relationship between mite abundance and any measure of body condition or fitness for either species or sex (also contrary to our predictions). Our results suggest that species biology and ecological context may influence mite abundance on hosts. However, this pattern does not extend to differential effects of mites on measures of host body condition or fitness.