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HierarcHical analysis of ampHibian Diversity in primary anD seconDary rain forests of central vietnam

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Abstract

We assessed species composition, diversity, and distribution patterns of rain forest amphibian assemblages in central Vietnam using a hierarchical analysis of diversity across study areas and different habitat types. We also plotted rarefaction curves to compare unequal assemblages of amphibians and compared the results with our field observations. We recorded 13,529 individuals of 100 species from Anura and Gymnophiona, comprising eight families and 33 genera. There was an overall similarity in species composition of 0.86 between two areas, A Luoi and Nam Dong, Thua Thien Hue Province, central Vietnam; however, the overall similarity in species composition between primary and secondary forests was only 0.03. Only 29 species (29%) of total recorded amphibians were found in both forest types. The relative abundance and species richness were significantly higher in primary forests than in secondary forests. In both areas, rarefaction curves revealed that the expected number of amphibian species in primary forests was significantly richer than that in secondary forests, suggesting that assemblages of amphibians in interior forest habitat may be more vulnerable to alterations caused by disturbances, such as logging. The lognormal model to predict the expected total number of species theoretically available for observation in the entire community (including the rare species not yet collected) indicated that the expected total number of species is approximately 105, with the number of unseen species predicted to fall below the veil line estimated at about 4.48.

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We used an experimental approach to investigate the effects of landscape composition on the initial dispersal success of juvenile amphibians. Larval amphibians—spotted salamander (Ambystoma maculatum), small-mouthed salamander (A. texanum), and American toad ( Bufo americanus )—were added to artificial pools in four dispersal arrays on forest edges. Each array consisted of a pool surrounded by a circular drift fence with pitfall traps and two 2.5 × 50 m enclosures (runs) extending into forest and old-field habitat. Juveniles captured at the circular fences were individually marked and released into either field or forest runs. We determined initial distance, initial rate, total distance, and net distance moved by juveniles in the field versus forest from recaptures in the runs. We also conducted 24-hour dehydration trials to compare the rates of evaporative water loss by spotted and small-mouthed salamanders in field and forest. Initial orientation of spotted salamanders and toads was significantly biased toward forest. Orientation of small-mouthed salamanders did not differ significantly from random expectations. The avoidance of open-canopy habitat by juvenile American toads in particular indicates that predictions of dispersal behavior based on adult habitat use may be misleading. Spotted salamanders moved almost four times farther and toads more than three times farther into the forest than into the field, and recapture rates of both species were much lower in the field. We attribute the lower recapture rates and shorter distances moved in the field to higher mortality due to desiccation or an abundance of predators. Juvenile spotted and small-mouthed salamanders experienced greater evaporative water loss in the field. Our data on movement behavior and dehydration rates suggest that old-field habitats offer greater landscape resistance to dispersing juveniles of some species. Thus, forest fragmentation is likely to reduce dispersal rates between local populations of these three species, with potentially negative consequences for population persistence in altered landscapes. Resumen: Utilizamos un enfoque experimental para investigar los efectos de la composición del paisaje sobre el éxito inicial de dispersión de anfibios juveniles. Colcamos larvas de anfibios (salamandras manchadas [Ambystoma maculatum] y A. texanum y sapo americano [Bufo americanus] ) en estanques artificiales en cuatro secuencias de dispersión en bordes de bosque. Cada secuencia consistió de un estanque rodeado por un cerco circular con trampas de fosa y dos encierros (corridas) de 2.5 × 50 m que se extendían hacia el hábitat de bosque y de campo viejo. Los juveniles capturados en los cercos circulares fueron marcados individualmente y liberados en las corridas de bosque o de campo. A partir de recapturas en las corridas, determinamos la distancia inicial, la tasa inicial, las distancia total y la distancia neta recorrida por juveniles en el campo versus el bosque. También realizamos pruebas de deshidratación de 24 horas para comparar las tasas de pérdida de agua por evaporación en salamandras en el campo y el bosque. La orientación inicial de Ambystoma maculatum y Bufo americanus estuvo significativamente sesgada hacia el bosque. La orientación inicial de A. texanum no fue significativamente diferente de las expectativas aleatorias. La evasión del hábitat abierto en particular por juveniles de sapo americano indica que las predicciones del comportamiento de dispersión basadas en el uso del hábitat por adultos pueden llevar a conclusiones erróneas. Las salamandras manchadas se movieron cuatro veces mas lejos y los sapos más de tres veces más lejos dentro del bosque que dentro del campo, y las tasas de recaptura de ambas especies fueron mucho menores en el campo. Atribuimos las bajas tasas de recaptura y las distancias menores a la mayor mortalidad debido a la desecación o a la abundancia de depredadores. Los juveniles de las dos especies de salamandras experimentaron mayor pérdida de agua por evaporación en los campos. Nuestros datos del comportamiento de movimiento y las tasas de deshidratación sugieren que los hábitats de campo viejo ofrecen mayor resistencia de paisaje para los juveniles dispersantes de algunas especies. Por tanto, es probable que la fragmentación de bosques reduce las tasas de dispersión entre poblaciones locales de estas tres especies, con consecuencias potencialmente negativas para la persistencia de la población en paisajes alterados.
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Recent warming has caused changes in species distribution and abundance, but the extent of the effects is unclear. Here we investigate whether such changes in highland forests at Monteverde, Costa Rica, are related to the increase in air temperatures that followed a step-like warming of tropical oceans in 1976 (refs4, 5). Twenty of 50 species of anurans (frogs and toads) in a 30-km2 study area, including the locally endemic golden toad (Bufo periglenes), disappeared following synchronous population crashes in 1987 (refs 6-8). Our results indicate that these crashes probably belong to a constellation of demographic changes that have altered communities of birds, reptiles and amphibians in the area and are linked to recent warming. The changes are all associated with patterns of dry-season mist frequency, which is negatively correlated with sea surface temperatures in the equatorial Pacific and has declined dramatically since the mid-1970s. The biological and climatic patterns suggest that atmospheric warming has raised the average altitude at the base of the orographic cloud bank, as predicted by the lifting-cloud-base hypothesis,.
Article
Large-scale amphibian conservation often relies on remotely sensed data to describe spatial patterns in occupancy. Commonly used data include ownership, forest type, soil class, and proximity of wetlands. We evaluated the influence of these variables on terrestrial amphibian occupancy in southern Michigan. We trapped amphibians using drift-fence arrays in 82 forest patches and estimated detection probabilities for all captured species. Ten species were captured, but only three were detected at levels suitable for occupancy modeling. We concluded that drift-fence arrays alone were not sufficient for adequately detecting the majority of terrestrial amphibians. The best detection models for American Toad (Anaxyrus americanus), Wood Frog (Lithobates sylvaticus), and Spring Peeper (Pseudacris crucifer) consistently included a positive relationship with temperature and precipitation. Detection-adjusted occupancy models for the three anurans indicated relatively high occupancy (>0.43). Ownership, forest type, and soil class did not occur in higher ranking occupancy models. Rather, the higher ranking models often included the distance to National Wetlands Inventory (NWI) wetland and sample year factors. However, these effects were highly variable and of no use in consistently predicting terrestrial amphibian occupancy. Our results substantiate the importance of incorporating detection probabilities into studies of amphibian habitat relationships and suggest that coarse descriptors of environmental conditions may not adequately portray factors important to terrestrial amphibian occupancy.
Article
Disentangling local and historical factors that determine species diversity patterns at multiple spatial scales is fundamental to elucidating processes that govern ecological communities. Here we investigated how environmental correlates may influence diversity at local and regional scales. Primarily utilizing published species lists, amphibian and reptile alpha and beta diversity were assessed at 17 well-surveyed sites distributed among ecoregions throughout Costa Rica. The degree to which regional species diversity patterns were related to environmental variables and geographic distance was determined using Canonical Correspondence Analysis and Mantel tests. Amphibian alpha diversity was highest in lowland Pacific sites (mean = 43.3 species) and lowest at the high elevation site (9 species). Reptile alpha diversity values were high for both lowland Atlantic (mean = 69.5 species) and lowland Pacific (mean = 67 species) sites and lowest for the high elevation site (8 species). We found high species turnover between local sites and ecoregions, demonstrating the importance of beta diversity in the determination of regional diversity. For both amphibians and reptiles, beta diversity was highest between the high-elevation site and all others, and lowest among lowland sites within the same ecoregion. The effect of geographic distance on beta diversity was minor. Ecologically significant climatic variables related to rain, temperature, sunshine and insolation were found to be important determinants of local and regional diversity for both amphibians and reptiles in Costa Rica.
Article
Effective monitoring programs are designed to track changes in the distribution, occurrence, and abundance of species. We developed an extension of Royle and Kéry's (2007) single species model to estimate simultaneously temporal changes in probabilities of detection, occupancy, colonization, extinction, and species turnover using data on calling anuran amphibians, collected from 2002 to 2006 in the Lower Mississippi Alluvial Valley of Louisiana, USA. During our 5-year study, estimates of occurrence probabilities declined for all 12 species detected. These declines occurred primarily in conjunction with variation in estimates of local extinction probabilities (cajun chorus frog [Pseudacris fouquettei], spring peeper [P. crucifer], northern cricket frog [Acris crepitans], Cope's gray treefrog [Hyla chrysoscelis], green treefrog [H. cinerea], squirrel treefrog [H. squirella], southern leopard frog [Lithobates sphenocephalus], bronze frog [L. clamitans], American bullfrog [L. catesbeianus], and Fowler's toad [Anaxyrus fowleri]). For 2 species (eastern narrow-mouthed toad [Gastrophryne carolinensis] and Gulf Coast toad [Incilius nebulifer]), declines in occupancy appeared to be a consequence of both increased local extinction and decreased colonization events. The eastern narrow-mouthed toad experienced a 2.5-fold increase in estimates of occupancy in 2004, possibly because of the high amount of rainfall received during that year, along with a decrease in extinction and increase in colonization of new sites between 2003 and 2004. Our model can be incorporated into monitoring programs to estimate simultaneously the occupancy dynamics for multiple species that show similar responses to ecological conditions. It will likely be an important asset for those monitoring programs that employ the same methods to sample assemblages of ecologically similar species, including those that are rare. By combining information from multiple species to decrease the variance on estimates of individual species, our results are advantageous compared to single-species models. This feature enables managers and researchers to use an entire community, rather than just one species, as an ecological indicator in monitoring programs. © 2011 The Wildlife Society.
Article
Over the last two decades, numerous studies have shown that alien predators contributed to amphibian population declines. Both experimental studies and correlative field surveys implicated alien species of fish, bullfrogs and crayfish as major contributors to amphibian population decline, and in some instances local extinction. Additional studies have demonstrated that alien predators also caused long-term changes in aquatic communities. Recent studies have examined the feasibility of removing alien predators, and provide some evidence that amphibian populations can recover. Applying information gained from past studies to the recovery of amphibian populations will be the challenge of future studies. International, national and local policies that regulate alien predators should be based largely on the body of scientific evidence already in the literature. Scientists need to be more involved with policy-makers to most effectively change laws that regulate alien predators.
Article
Ten species of small vertebrates that occurred together in leaf litter of the Amazon tropical rainforest were studied during the dry season in Rondônia, western Brazil. This vertebrate assemblage consisted of two diurnal lizards, two diurnal frogs, one nocturnal frog, and five frog species that are active by day and night. All species feed on small invertebrates, primarily insects and arachnids. There are significant differences in prey types and sizes eaten, and overlap values among species are relatively low. Niche breadths vary from unity (extremely low) to high (> 9.0). A pseudocommunity analysis using two different algorithms indicated that structure exists in the prey resource matrix of the vertebrate assemblage. The zero structure of the community matrix contributes to the observed structure, i.e. prey items not taken by certain species are important in maintaining structure. Several well-defined feeding guilds are apparent, and the feeding guilds do not appear to follow taxonomic lines.
Article
In community ecology, contrasting theories suggest that the distribution and abundance of species, and thus the composition of assemblages, are influenced by i) environmental gradients, or ii) contagious biotic processes such as predation, competition, dispersal and disease. In the former case, sites with similar environments would tend to support similar assemblages, while in the latter, geographically proximate sites would tend to support more similar assemblages than widely separated sites. I investigated the relative influence of environmental variables and spatial position on the composition of frog assemblages at forest streams in sub-tropical eastern Australia using redundancy analysis (RDA) and partial RDA. Data on the maximum abundance of the frog species at 65 survey sites were transformed such that RDA would yield the Hellinger distance between sites. The following analysis identified 11 environmental variables that explained 45% of the variation in the abundance of species at the survey sites (the species matrix), as a proportion of total variance. The geographic co-ordinates of the survey sites accounted for 12%, while the environmental and spatial variables combined accounted for 47% of the variation in the species matrix. Partial redundancy analysis indicated that of the explained variation, 74% was purely environmental, 5% was purely spatial and 21% was spatial environmental variation. This study is the first to quantify the relative influence of environmental and spatial variables on the composition of amphibian assemblages. It provides support for both the environmental control model and the biotic control model of species’ distributions and assemblage composition, although environmental variables appear to have the greater effect at this scale of analysis.
Article
We present data on sample richness, relative abundance, and community structure of a leaf litter amphibian assemblage from globally important miombo-mopane woodlands characteristic of western Tanzania. We describe patterns of diversity across major habitat types and between different seasons from an annual pitfall-trapping campaign. We recorded 28 species of amphibians, which is significantly higher than existing richness estimates for other miombo woodland sites elsewhere in sub-Saharan Africa. We found that cultivation of native habitat reduces frog diversity, a conclusion that has important implications in light of the rapid conversion of miombo woodland for agriculture and fuel-wood across much of southern and central Africa. Many species showed strongly ansynchronized patterns of seasonality in relative abundance, which has significant implications for the establishment of successful monitoring programs and biodiversity surveys. These conclusions emphasize the importance of stratified long-term sampling in biodiversity studies and demonstrate that superficial levels of sampling effort can lead to erroneous conclusions regarding patterns of diversity in amphibian communities. The relatively poor focus on herpetofaunal research in African miombo-mopane woodland is out of proportion to its ecological and conservation significance.
Article
Relationships between diurnal, terrestrial birds, vegetation and environmental attributes were investigated in the open Eucalyptus forests of south east Australia. Of the 83 bird species recorded, the probability of occurrence of the Yellow-faced Honeyeater, Superb Fairy-wren, Golden Whistler, Fan-tailed Cuckoo and Shining Bronze-Cuckoo was modelled. The Yellow-faced Honeyeater was the only taxon modelled whose distribution could be best explained using climatic variables. The other four species were related to habitat features at the site level. The models developed for these species were specific to each data set used and are unlikely to be appropriate for other data collected at other times of the year. Statistical modelling was used to investigate the response of bird assemblages to environmental and biological regimes. The models reflect differences in the bird species composition of sites in relation to elevation and soil moisture/nutrient gradients across the region. Birds favouring higher elevations were separated from birds favouring mid and lower elevations. Birds favouring wetter environments were separated from those using drier habitats although these species were also present in the wetter environments. Differences in the distribution of birds related to elevation were explained by the environmental attributes minimum temperature of the coldest month and mean precipitation of the driest quarter which also affect vegetation formations and associations across the study region. Vegetation floristics and topographic position, which can reflect the moisture and nutrient status of sites, were important in characterising the specific habitat requirements of a number of birds. Physical and biological regimes were significantly related to bird species richness. Species richness was found to be greatest in moist forests found at mid and low elevations throughout the year. A stratified field survey was successful in recording the majority of forest birds that can occur in the eucalypt forests of south east Australia but sufficient data for reliably modelling the distribution and habitat quality of birds were obtained for only about 10% of the birds recorded. Even when data from two census periods were aggregated, data were sufficient for only 27% of the total number of birds recorded to develop a habitat model. Suggested modifications to the survey design based on the data and analyses used in this study, are discussed in terms of the intended purpose of the field survey.
Article
Many indices have been proposed for measuring species evenness in ecological communities, but there is no consensus on which are best. We assemble criteria for an appropriate index for the evenness of a biological sample. The most important criterion is that evenness should be independent of species richness. Twelve previously proposed indices and variants are considered, and two apparently new indices. Four indices are recommended as joint best buys: A) If symmetry between minor and abundant species is not important, or if it is required that the index be less affected by minor species: 1) If it is essential that the index be able to reach a minimum of zero with any particular number of species, or if the shape of the index response to an evenness gradient is important: E1/D (based on a common form of Simpson's index). 2) If good mid-range behaviour is desired: E′ (proposed by Camargo). B) If equal sensitivity to minor and abundant species is required: 1) If the shape of the index response to an evenness gradient is not important, the clear winner is: EQ (a new index). 2) If the shape is important: Evar (another new index). The overall recommendation for general use is Evar.
Article
In this paper a methodology is presented for measuring diversity based on rarefaction of actual samples. By the use of this technique, a within-habitat analysis was made of the bivalve and polychaete components of soft-bottom marine faunas which differed in latitude, depth, temperature, and salinity. The resulting diversity values were highly correlated with the physical stability and past history of these environments. A stability-time hypothesis was invoked to fit these findings, and, with this hypothesis, predictions were made about the diversities present in certain other environments as yet unstudied. The two types of diversity, based on numerical percentage composition and on number of species, were compared and shown to be poorly correlated with each other. Our data indicated that species number is the more valid diversity measurement. The rarefaction methodology was compared with a number of diversity indexes using identical data. Many of these indexes were markedly influenced by sample size. Good...
Article
Species richness, Shannon information, and Simpson diversity are the three most commonly used nonparametric measures of species diversity. The sampling bias and variance of these measures differ greatly. Species richness may be seriously underestimated for even very large samples from a speciose community. The bias in species richness and Shannon information depend on unknown parameters of the species abundance distribution. An unbiased estimator exists only for Simpson diversity. Each of these measures is concave, so that the total diversity in a pooled set of communities exceeds (or equals) the average diversity within communities. The total diversity in a set of communities can therefore be partitioned into positive, additive components within and among communities, corresponding to alpha- and beta-diversity. Partitioning Simpson diversity corresponds to an analysis of variance. The proportion of the total diversity found within communities provides a natural measure of similarity among multiple communities. The expected similarity among multiple random samples from the same community depends on the number of samples and on the underlying measure of diversity.
Article
Part 1. It is shown that in a large collection of Lepidoptera captured in Malaya the frequency of the number of species represented by different numbers of individuals fitted somewhat closely to a hyperbola type of curve, so long as only the rarer species were considered. The data for the commoner species was not so strictly `randomized', but the whole series could be closely fitted by a series of the logarithmic type as described by Fisher in Part 3. Other data for random collections of insects in the field were also shown to fit fairly well to this series. Part 2. Extensive data on the capture of about 1500 Macrolepidoptera of about 240 species in a light-trap at Harpenden is analysed in relation to Fisher's mathematical theory and is shown to fit extremely closely to the calculations. The calculations are applied first to the frequency of occurrence of species represented by different numbers of individuals--and secondly to the number of species in samples of different sizes from the same population. The parameter ` alpha ', which it is suggested should be called the `index of diversity', is shown to have a regular seasonal change in the case of the Macrolepidoptera in the trap. In addition, samples from two traps which overlooked somewhat different vegetation are shown to have ` alpha ' values which are significantly different. It is shown that, provided the samples are not small, ` alpha ' is the increase in the number of species obtained by increasing the size of a sample by e (2.718). A diagram is given (Fig. 8) from which any one of the values, total number of species, total number of individuals and index of diversity (alpha), can be obtained approximately if the other two are known. The standard error of alpha is also indicated on the same diagram. Part 3. A theoretical distribution is developed which appears to be suitable for the frequencies with which different species occur in a random collection, in the common case in which many species are so rare that their chance of inclusion is small. The relationships of the new distribution with the negative binomial and the Poisson series are established. Numerical processes are exhibited for fitting the series to observations containing given numbers of species and individuals, and for estimating the parameter alpha representing the richness in species of the material sampled; secondly, for calculating the standard error of alpha, and thirdly, for testing whether the series exhibits a significant deviation from the limiting form used. Special tables are presented for facilitating these calculations.
Article
The recent literature on species diversity contains many semantic, conceptual, and technical problems. It is suggested that, as a result of these problems, species diversity has become a meaningless concept, that the term be abandoned, and that ecologists take a more critical approach to species-number relations and rely less on information theoretic and other analogies. As multispecific collections of organisms possess numerous statistical properties which conform to the conventional criteria for diversity indices, such collections are not intrinsically arrangeable in linear order along some diversity scale. Several such properties or "species composition parameters" having straightforward biological interpretations are presented as alternatives to the diversity approach. The two most basic of these are simply Δ1=[NN1][1Σi(NiN)2]\Delta_1= [\frac{N}{N-1}][^1-\Sigma_i (\frac{N_i} {N})^2] =the proportion of potential interindividual encounters which is interspecific (as opposed to intraspecific), assuming every individual in the collection can encounter all other individuals, and E(Sn)=iΣ[1((NNin))((Nn))]E(S_n)= ^\Sigma_i [1-\frac{(\binom{N-N_i}{n})}{(\binom{N}{n})}] =the expected number of species in a sample of n individuals selected at random from a collection containing N individuals, S species, and Ni individuals in the ith species.
Article
Consisting of more than six thousand species, amphibians are more diverse than mammals and are found on every continent save Antarctica. Despite the abundance and diversity of these animals, many aspects of the biology of amphibians remain unstudied or misunderstood. The Ecology and Behavior of Amphibians aims to fill this gap in the literature on this remarkable taxon. It is a celebration of the diversity of amphibian life and the ecological and behavioral adaptations that have made it a successful component of terrestrial and aquatic ecosystems. Synthesizing seventy years of research on amphibian biology, Kentwood D. Wells addresses all major areas of inquiry, including phylogeny, classification, and morphology; aspects of physiological ecology such as water and temperature relations, respiration, metabolism, and energetics; movements and orientation; communication and social behavior; reproduction and parental care; ecology and behavior of amphibian larvae and ecological aspects of metamorphosis; ecological impact of predation on amphibian populations and antipredator defenses; and aspects of amphibian community ecology. With an eye towards modern concerns, The Ecology and Behavior of Amphibians concludes with a chapter devoted to amphibian conservation. An unprecedented scholarly contribution to amphibian biology, this book is eagerly anticipated among specialists.
Article
Conservationists are far from able to assist all species under threat, if only for lack of funding. This places a premium on priorities: how can we support the most species at the least cost? One way is to identify 'biodiversity hotspots' where exceptional concentrations of endemic species are undergoing exceptional loss of habitat. As many as 44% of all species of vascular plants and 35% of all species in four vertebrate groups are confined to 25 hotspots comprising only 1.4% of the land surface of the Earth. This opens the way for a 'silver bullet' strategy on the part of conservation planners, focusing on these hotspots in proportion to their share of the world's species at risk.
Ecological Methodology
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Amphibian diversity in central Vietnam and the distribution pattern and natural history of Granular Spiny Frogs Quasipaa verrucospinosa (Anura: Dicroglossidae)
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