Comparison of Two Techniques for Surveying Headwater Stream Amphibians

Abstract: We compared rubble-rousing versus light-touch stream amphibian survey techniques in multiple 1-m plots across 10 streams in southwest Washington, USA. Specifically, we wanted to determine if light-touch surveys provide unbiased estimates of abundance (i.e., provide counts correlated with rubble-rousing counts) and which method would provide more cost-effective presence or absence information. Rubble-rousing, a common technique for surveying stream-associated amphibians in the Pacific Northwest, took 12 times as long as light-touch to apply. Abundance estimates and standard errors for rubble-rousing were consistently higher than those for light-touch for all life stages for the coastal tailed frog (Ascaphus truei) and Columbia torrent salamander (Rhyacotriton kezeri). Except for eggs, light-touch detected all life stages found during rubble-rousing. For frogs, only some rubble-rousing abundance estimates, mostly involving second-year larvae, were highly correlated with their light-touch counterparts, whereas for salamanders, similar comparisons generated high correlations across most life stages. Correlations between methods were consistently greater for salamanders than for frogs. However the smaller tailed frog sample sizes and the cryptozoic nature of some life stages may have contributed to this pattern. Depending on the degree to which researchers can tolerate false-negative error rates, light-touch may prove less costly than rubble-rousing for detecting species presence. For the cost of obtaining one rubble-rousing sample, many light-touch samples can be used across a range of habitats for detecting species patchily distributed.     

Basin-Scale Surveys of Stream-Associated Amphibians in Intensively Managed Forests

Abstract: Conservation and management of native species on landscapes managed for intensive wood production represents an ongoing challenge to forest managers. Previous research suggests that impacts of forest practices on stream-associated amphibians (SAA; giant [Dicamptodon spp.], torrent [Rhyacotriton spp.], and plethodontid [Plethodon spp.] salamanders and coastal tailed frogs [Ascaphus truei]) in Oregon and Washington, USA, vary spatially and temporally as a result of biotic and abiotic factors, some of which can be influenced by management treatments. Although individual harvest units can encompass multiple stream reaches and entire second-order basins, nearly all published research studies used stream reaches of various lengths as sample units. To address this discrepancy between research and operational scales, we sampled first-, second-, and third-order streams in 70 randomly selected third-order basins in Oregon and Washington in 2007 and 2008 to estimate detection and occupancy parameters for SAA and to develop basin-level density estimates for different species and genera. We estimated occupancy probabilities of 0.99 (95% CL = 0.96–1.00) for torrent and giant salamanders, 0.93 (95% CL = 0.76–0.92) for Dunn's salamanders (Plethodon dunni), and 0.60 (95% CL = 0.46–0.72) for tailed frogs. Our estimates can be compared with estimates for unmanaged third-order basins in Oregon and Washington to provide a relative measure of potential impacts of forest management on these taxa. In addition, our estimates provide baseline information with which to assess potential effects of future environmental changes on the 4 genera.     

Topographic microclimates drive microhabitat associations at the range margin of a butterfly

The habitat associations of individuals underpin the dynamics of species distributions. Broad‐scale gradients in climate can alter habitat associations across species’ geographic ranges, but topographic heterogeneity creates local microclimates which could generate variation in habitat use at finer spatial scales. We examined the selection of microhabitats for egg‐laying by populations of a thermally‐constrained butterfly, the skipper Hesperia comma, across 16 sites with different regional temperatures and topographic microclimates. Using models of thermal microclimate, we examined how the association between eggs and warm bare ground microhabitats varied with ambient temperature, and predicted bare ground associations in 287 existing H. comma populations, to investigate the relative impacts of regional temperatures and topographic microclimates on microhabitat use. Eggs were most strongly associated with bare ground in relatively cool sites, indicating climate‐driven changes in microhabitat use. The majority of temperature variation between study sites was attributable to topographic microclimates rather than regional temperature differences, such that changes in microhabitat associations occurred principally between north‐ and south‐facing slopes within the same region. Predicted microhabitat associations across the UK distribution of H. comma showed that, due to the large temperature differences generated by topography, most of the between‐population variation in microhabitat use occurs locally within 5 km grid squares, with a smaller proportion occurring at a regional level between 5 km squares. Our findings show how microclimatic variation generated by topography alters the habitat associations of populations at fine spatial scales, suggesting that microclimate‐driven changes in habitat suitability could shape species’ distribution dynamics and their responses to environmental change.     

Rangewide landscape genetics of an endemic Pacific northwestern salamander

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.     

The area-independent effects of habitat complexity on biodiversity vary between regions

Potential explanatory variables often co‐vary in studies of species richness. Where topography varies within a survey it is difficult to separate area and habitat‐diversity effects. Topographically complex surfaces may contain more species due to increased habitat diversity or as a result of increased area per se. Fractal geometry can be used to adjust species richness estimates to control for increases in area on complex surfaces. Application of fractal techniques to a survey of rocky shores demonstrated an unambiguous area‐independent effect of topography on species richness in the Isle of Man. In contrast, variation in species richness in south‐west England reflected surface availability alone. Multivariate tests and variation in limpet abundances also demonstrated regional variation in the area‐independent effects of topography. Community composition did not vary with increasing surface complexity in south‐west England. These results suggest large‐scale gradients in the effects of heterogeneity on community processes or demography.     

Shallow environmental gradients put inland species at risk: Insights and implications from predicting future distributions of Eucalyptus species in South Western Australia

As climate changes, tree decline in Mediterranean‐type ecosystems is increasing worldwide, often due to decreased effective precipitation and increased drought and heat stress, and has recently been observed in coastal species of the iconic Eucalyptus (Myrtaceae) genus in the biodiversity hotspot of south‐west Western Australia. To investigate how this drought‐related decline is likely to continue in the future, we used species distribution modelling techniques to generate broad‐scale predictions of future distribution patterns under three distinct projected climate change scenarios. In a moderate climate change scenario, suitable habitat for all species was predicted to decrease by, on average, 73% by the year 2100, with most receding into southern areas of their current distribution. Although the most severe Eucalyptus declines in south‐west Western Australia have been observed in near‐coastal regions, our predictions suggest that inland species are at greater risk from climate change, with six inland species predicted to lose 95% of their suitable habitat in a moderate change scenario. This is due to the shallow environmental gradients of inland regions causing larger spatial shifts of environmental envelopes, which is likely to be relevant in many regions of the world. The knowledge gained suggests that future research and conservation efforts in south‐west Western Australia and elsewhere should avoid focussing disproportionately on coastal regions for reasons of convenience and proximity to population centres, and properly address the inland region where the biggest future impacts may occur.     

Characterization of an Atlantic salmon Salmo salar stream at the southern limit of its eastern Atlantic distribution

Ten reaches of an Atlantic stream located in north‐west Spain were sampled intensively during one summer to characterize the conditions where Atlantic salmon Salmo salar have been re‐introduced along the stream. Fish species richness and diversity showed a downstream increase, which was mainly attributed to the higher number of cyprinid species found in the lower reaches. Moreover, except for brown trout Salmo trutta that appeared to be the most ubiquitous species, the densities of the other species was higher in the lower than in the upper stream reaches. Redundancy analysis showed that the pattern of fish assemblages observed along the studied stream was mainly related to the expected gradient observed in the levels of dissolved oxygen, discharge and mean current speed. There was a significant differentiation between midstream and downstream reaches, both in terms of the composition of their fish assemblages and the freshwater habitat. This study emphasizes the importance of describing the variations in fish assemblages and habitat characteristics along a river to explore its relation to potential changes in the survival of fish populations. In particular, the development of habitat–fish relationships may be a useful tool for water managers to assess the effects of development and restoration projects on the habitat of S. salar.     

Contrasting responses of Peromyscus mice of Yosemite National Park to recent climate change

Accurately predicting the response of species to climate change is crucial for the preservation of contemporary species diversity. In the current study, we analyze the response of two congeneric small mammal species (Peromyscus maniculatus and Peromyscus truei) to recent climate change in the region of Yosemite National Park (California, USA). The generalist P. maniculatus did not change its distribution in response to climate change while the specialist P. truei substantially changed its geographic and elevational distribution in the region, expanding into Yosemite. Using molecular genetic techniques we found that a cryptic geographic shift in genetic variation may have occurred within the geographically stable P. maniculatus distribution. Using a combination of morphometric and molecular genetic techniques we confirmed that a P. truei subspecies previously identified as a habitat specialist expanded into new habitat types, suggesting that this subspecies is not in fact a habitat specialist. Instead, we propose that the range of this subspecies is instead limited by climatic variables currently varying in response to contemporary climate change. These results underscore the importance of verifying the natural‐history‐based assumptions used to develop predictive models of species' response to climate change.     

Microhabitat relationships among five lizard species associated with granite outcrops in fragmented agricultural landscapes of south‐eastern Australia

A fundamental part of developing effective biodiversity conservation is to understand what factors affect the distribution and abundance of particular species. However, there is a paucity of data on ecological requirements and habitat relationships for many species, especially for groups such as reptiles. Furthermore, it is not clear whether habitat relationships for particular species in a given environment are transferable to other environments within their geographical range. This has implications for the type of ‘landscape model’ used to guide management decisions in different environments worldwide. To test the hypothesis that species‐specific habitat relationships are transferable to other environments, we present microhabitat models for five common lizard species from a poorly studied habitat – insular granite outcrops, and then compared these relationships with studies from other environments in south‐eastern Australia. We recorded twelve species from five families, representing 699 individuals, from 44 outcrops in the south‐west slopes of New South Wales. Five lizard species were abundant and accounted for 95% of all observations: Egernia striolata, Ctenotus robustus, Cryptoblepharus carnabyi, Morethia boulengeri and Carlia tetradactyla (Scincidae). Linear regression modelling revealed suites of different variables related to the abundance patterns of individual species, some of which were broadly congruent with those measured for each species in other environments. However, additional variables, particular to rocky environments, were found to relate to reptile abundance in this environment. This finding means that species' habitat relationships in one habitat may not be readily transferable to other environments, even those relatively close by. Based on these data, management decisions targeting reptile conservation in agricultural landscapes, which contain rocky outcrops, will be best guided by landscape models that not only recognize gradients in habitat suitability, but are also flexible enough to incorporate intraspecies habitat variability.     

Three new species of Callulina (Amphibia: Anura: Brevicipitidae) highlight local endemism and conservation plight of Africa's Eastern Arc forests

Material ascribed to the genus Callulina from north‐east Tanzania and south‐east Kenya is assessed. Three new species of Callulina are described from the North ( Callulina laphami sp. nov. ) and South ( Callulina shengena sp. nov. and Callulina stanleyi sp. nov. ) Pare Mountains in Tanzania. The species are diagnosed based on morphological, acoustic, and molecular data. A new key to the species of Callulina is provided. Based on an interpretation of the International Union for Conservation of Nature (IUCN) red list, we suggest that the three species will qualify as critically endangered, because of their small distributions and the ongoing threat to their habitat. We reveal the high local endemism of Callulina in the northern part of the Eastern Arc Mountains, with each species restricted to no more than one mountain (fragment) block. This high local endemism in Callulina is probably widespread across the Eastern Arc, raising further conservation concern for this group of amphibians. Based on new molecular phylogenetic data for Callulina, we discuss biogeographical relationships among north‐east Tanzanian mountains, and evolutionary patterns in Eastern Arc breviciptids. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 496–514.     

Stream amphibians as metrics of critical biological thresholds in the Pacific Northwest, U.S.A.: a response to Kroll et al.

1. Kroll, Hayes & MacCracken (in press) Concerns regarding the use of amphibians as metrics of critical biological thresholds: a comment on Welsh and Hodgson 2008 . Freshwater Biology, criticised our paper [ Welsh & Hodgson (2008) Amphibians as metrics of critical biological thresholds in forested headwater streams of the Pacific Northwest. Freshwater Biology, 53 , 1470–1488] proposing the use of headwater stream amphibians as metrics of stream status in the Pacific Northwest (PNW). They argued that our analysis of previously published data reflected circular reasoning because we reached the same conclusions as the earlier studies. In fact, we conducted a meta‐analysis to address new questions about the optimum values and thresholds (based on animal densities) for abiotic stream attributes that were found to be important to these amphibians in earlier studies. This is analogous to determining blood pressure thresholds or fat‐to‐weight ratios that facilitate predicting human health based on meta‐analyses of earlier data from studies that found significant correlations between these variables and relative health. 2. Kroll et al. argued that we should not make inference to environmental conditions across the PNW from data collected in California. We collected data from northern California and southern Oregon, the southern extent of the PNW. We made inference to the Klamath‐Siskiyou and North Coast bioregions, and argued that available research on these headwater species indicates that our results have the potential to be applied throughout the PNW with minimal regional adjustments. 3. Kroll et al. contended that we need reproductive success, survival estimates and density estimates, corrected for detection probabilities, to establish relationships between animal density and stream attributes. Reproductive success and survival estimates are important for demographic modelling and life tables, but they are not necessary to demonstrate meaningful relationships with abiotic conditions. Both corrected occupancy estimates and individual detection probabilities are unnecessary, and take multiple sampling efforts per site, or onerous mark release and re‐capture studies, respectively, to determine accurately. 4. Kroll et al. questioned the use of stream amphibians as a surrogate for measuring physical parameters, such as water temperature, claiming that measuring the physical parameters directly is more efficient. Here they misinterpreted the main point of our paper: stream organisms are integrators of what happens in a catchment, and carefully selected species can serve as surrogates for the biotic community and the relative condition of the network environment. 5. Kroll et al. claimed that we demonstrated weak inferences regarding ecosystem processes. We argue that by relating densities of stream amphibians with changes along abiotic environmental gradients that are commonly affected by anthropogenic activities, we are establishing biological links to gradients that represent important ecosystem processes and identifying biometrics that can be used to quantify the status (health) of these gradients.     

Patterns of geophyte diversity and storage organ size in the winter-rainfall region of southern Africa

The winter‐rainfall region of southern Africa, covered largely by the fynbos and succulent karoo biomes, harbours the world's greatest concentration of geophyte species. Species diversity is greatest in the south‐west, where more than 500 species co‐occur in one quarter‐degree square; in the south‐east the values are generally around 100, and in the arid north‐west, always less than 50 (more often less than 10). In at least three species‐rich genera (Moraea, Eriospermum and Oxalis), the size of storage organs (bulbs, corms, tubers) varies inversely, with the largest average values occurring in the species‐poorer areas — both in the north‐western, and in the south‐eastern parts of the region. This negative correlation between average storage organ size and species diversity is, however, only observed at relatively large spatial scales, which suggests that there is no direct relationship between storage organ size and species diversity. More likely, both these measures are driven by winter rainfall amount and reliability, both of which peak in the south‐western Cape. We suggest that reliable winter rainfall makes large storage organs unnecessary and depresses extinction rates, thus leading to the accumulation of species.     

Magnetic Compass Orientation in Larval Iberian Green Frogs, Pelophylax Perezi

Experiments were carried out to investigate whether Iberian green frog tadpoles Pelophylax perezi (formerly Rana perezi) are able of using the geomagnetic field for y‐axis orientation (i.e. orientation toward and away from shore). Tadpoles were trained outdoor for 5 d, in two different training configurations: (i) a training tank aligned along the magnetic north–south axis, with shore facing south, and (ii) a training tank aligned along the magnetic east–west axis, with shore located east, and similar to the shore–deep water axis (‘y‐axis’) found in their home stream, which flows from south to north. After training, tadpoles were individually tested for magnetic orientation in a water‐filled circular outdoor arena surrounded by a pair of orthogonally aligned cube‐surface‐coils used to alter the alignment of the earth's magnetic field. Tadpoles held in the east–west training tank oriented towards shore, indicating that they were able to distinguish between the shoreward and waterward direction along the y‐axis. Tadpoles trained in the tank that was aligned along the north–south axis showed bimodal magnetic compass orientation along the shore–deep water magnetic axis. These findings provide evidence for the use of magnetic compass cues for y‐axis orientation by P. perezi tadpoles.     

Marine reserves indirectly affect fine‐scale habitat associations,but not overall densities,of small benthic fishes

Many large, fishery‐targeted predatory species have attained very high relative densities as a direct result of protection by no‐take marine reserves. Indirect effects, via interactions with targeted species, may also occur for species that are not themselves targeted by fishing. In some temperate rocky reef ecosystems, indirect effects have caused profound changes in community structure, notably the restoration of predator–urchin–macroalgae trophic cascades. Yet, indirect effects on small benthic reef fishes remain poorly understood, perhaps because of behavioral associations with complex, refuge‐providing habitats. Few, if any, studies have evaluated any potential effects of marine reserves on habitat associations in small benthic fishes. We surveyed densities of small benthic fishes, including some endemic species of triplefin (Tripterygiidae), along with fine‐scale habitat features in kelp forests on rocky reefs in and around multiple marine reserves in northern New Zealand over 3 years. Bayesian generalized linear mixed models were used to evaluate evidence for (1) main effects of marine reserve protection, (2) associations with habitat gradients, including complexity, and (3) differences in habitat associations inside versus outside reserves. No evidence of overall main effects of marine reserves on species richness or densities of fishes was found. Both richness and densities showed strong associations with gradients in habitat features, particularly habitat complexity. In addition, some species exhibited reserve‐by‐habitat interactions, having different associations with habitat gradients inside versus outside marine reserves. Two species (Ruanoho whero and Forsterygion flavonigrum) showed stronger positive associations with habitat complexity inside reserves. These results are consistent with the presence of a behavioral risk effect, whereby prey fishes are more strongly attracted to habitats that provide refuge from predation in areas where predators are more abundant. This work highlights the importance of habitat structure and the potential for fishing to affect behavioral interactions and the interspecific dynamic attributes of community structure beyond simple predator–prey consumption and archetypal trophic cascades.     

Plant—herbivore interactions in streams near Mount St Helens

1. In four separate field experiments near Mount St Helens (Washington, U.S.A.) during 1986, the grazing effects of two large benthic herbivores, tadpoles of the tailed frog Ascaphus truei and larvae of the caddisfly Dicosmoecus gilvipes, were investigated using streamside channels and in-stream manipulations. In the experimental channels, abundances of periphyton and small benthic invertebrates declined significantly with increasing density of these larger herbivores. 2. In eleven small, high-gradient streams affected to varying degrees by the May 1980 eruption, in-stream platforms were used to reduce grazing by A, truei tadpoles on tile substrates. Single platforms erected in each tributary and compared to grazed controls revealed only minor grazing effects, and no significant differences among streams varying in disturbance intensity (and, consequently, tadpole density). However, results probably were confounded by high variability among streams in factors other than tadpole abundance. 3. Grazing effects were further examined in two unshaded streams with different tadpole densities, using five platforms per stream. In the stream with five tadpoles m^?2, grazing reduced periphyton biomass by 98% and chlorophyll a by 82%. In the stream lacking tadpoles, no significant grazing effects were revealed. Low algal abundance on both platforms and controls, and high invertebrate density in that stream (c. 30000m^?2) suggests that grazing by small, vagile invertebrates was approximately equivalent to that of tadpoles. 4. The influence of large benthic herbivores on algal and invertebrate communities in streams of Mount St Helens can be important, but reponses vary spatially in relation to stream disturbance history, local environmental factors, and herbivore distributional patterns and abundance.     

Distribution of saltmarsh plant communities associated with environmental factors along a latitudinal gradient on the south‐west Atlantic coast

Aim To produce an inventory of south‐west Atlantic saltmarshes (from latitude 31°48′ S to 43°20′ S) using remotely sensed images and field sampling; to quantify their total area; to describe the biogeographical variation of the main habitats characterized by dominant vascular plants, in relation to major environmental factors; to test the hypothesis of predominance of the reversal pattern in plant distribution (sedges and grasses dominate the lower, regularly inundated zones, while the upper zones are occupied by more halophytic species) previously described; and to compare these south‐west Atlantic saltmarshes with others world‐wide. Location South‐western Atlantic saltmarshes Methods Field samples of dominant emergent plant species positioned by the global positioning system (GPS) were obtained from most coastal saltmarshes (14) between southern Brazil and northern Patagonia, Argentina. Landsat satellite images were obtained and coastal saltmarsh habitats were quantified by supervised classification, utilizing points gathered in the field. Results Three main plant species dominated the low and middle intertidal saltmarsh, Spartina alterniflora Loesel., Spartina densiflora Brong. and Sarcocornia perennis (P. Mill.) A.J. Scott. The total area of the studied coastal saltmarshes was 2133 km², comprising 380 km² of Sp. alterniflora marsh, 366 km² of Sp. densiflora marsh, 746 km² of Sar. perennis marsh and 641 km² of brackish marsh (dominated by Juncus acutus L., Juncus kraussii Hochst., Scirpus maritimus L., Scirpus americanus Pers. and Phragmites australis (Cav.) Trin.). Cluster analysis showed three habitat types: saltmarshes dominated by (1) Sp. densiflora and brackish species,(2) Sp. alterniflora and Sar. perennis and (3) Sp.densiflora only. The analysis of abiotic variables showed significant differences between groups of habitats and coordinated gradients of the abiotic variables. The south‐west Atlantic coast showed decreasing mean annual rainfall (1200 to 196 mm) and increasing mean tidal amplitude (< 0.5 to > 2.5 m) from latitude 31° to 43°. Main conclusions South‐west Atlantic saltmarshes are globally important by virtue of their total extent. Remote sensing showed that the reversal pattern in plant distribution is not widespread. Indeed, south‐west Atlantic saltmarshes are better characterized by the presence of the halophytic genera Spartina and Sarcocornia. Our results support the interpretation that south‐west Atlantic saltmarshes constitute a class of temperate type (sensu Adam, 1990 ) with transitional characteristics between Australasian–South African saltmarshes and west Atlantic saltmarshes.     

706. BETULA ASHBURNERI

A high altitude shrub birch from Bhutan, south‐east Tibet, north‐west Yunnan and western Sichuan, which has been referred to the tetraploid Betula utilis, is shown to be a distinct diploid species, described here as Betula ashburneri McAllister & Rushforth.     

The ecological significance of the zooplanktivores, snipefish Macroramphosus spp. and boarfish Capros aper, in the food web of the south-east North Atlantic

The diets of slender snipefish Macroramphosus gracilis, longspine snipefish Macroramphosus scolopax and boarfish Capros aper, three very abundant species on the Portuguese coast, were studied from samples collected between July 2002 and April 2003. Variations in the diet with fish size, season and area, as well as diet overlap and diversity, are explored in this study. The diets of slender snipefish and boarfish were mainly composed of copepods, with the main prey being Temora spp. and Calanus spp., respectively. Mysid shrimps were the most important food item in the diet of longspine snipefish. During the summer season, when the availability of different prey items was highest, the two species of snipefish and the boarfish fed on different prey. Temora spp. were the most important prey in the stomachs of slender snipefish in the summer on the south‐west coast, while Calanus spp. started appearing in the stomachs of boarfish in the spring in the north, increasing their abundance in the summer on the south and south‐west coasts. The abundance of mysids appeared to increase in the autumn on the south‐west coast, being at that time an important food item for both longspine snipefish and boarfish. For slender snipefish and boarfish, the differences in stomach species diversity were explained firstly by the season and then by the area and fish size. For longspine snipefish, the area did not explain the species diversity variability, season being the first variable determining the differences. Of all three species, slender snipefish was the one with highest diversity of stomach contents, particularly in spring and summer on the north and south‐west coasts. Diet overlap between species was very low. No predation on eggs and larval stages of fishes was found for any of these fish species. This work is the first to address the diets of snipefish species and boarfish in the south‐east North Atlantic, in a large spatio‐temporal coverage. These species are important prey for many commercial species on the Portuguese coast and, given their abundance, may have a great impact on zooplankton communities, thus assuming a pivotal position in marine food webs.     

Squamate richness in the Brazilian Cerrado and its environmental–climatic associations

We investigate patterns of species richness of squamates (lizards, snakes, and amphisbaenians) in the Brazilian Cerrado, identifying areas of particularly high richness, and testing predictions of large‐scale richness hypotheses by analysing the relationship between species richness and environmental climatic variables. We used point localities from museum collections to produce maps of the predicted distributions for 237 Cerrado squamate species, using niche‐modelling techniques. We superimposed distributions of all species on a composite map, depicting richness across the ecosystem. Then, we performed a multiple regression analysis using eigenvector‐based spatial filtering (Principal Coordinate of Neighbour Matrices) to assess environmental–climatic variables that are best predictors of species richness. We found that the environmental–climatic and spatial filters multiple regression model explained 78% of the variation in Cerrado squamate richness (r² = 0.78; F = 32.66; P < 0.01). Best predictors of species richness were: annual precipitation, precipitation seasonality, altitude, net primary productivity, and precipitation during the driest quarter. A model selection approach revealed that several mechanisms related to the different diversity hypothesis might work together to explain richness variation in the Cerrado. Areas of higher species richness in Cerrado were located mainly in the south‐west, north, extreme east, and scattered areas in the north‐west portions of the biome. Partitioning of energy among species, habitat differentiation, and tolerance to variable environments may be the primary ecological factors determining variation in squamate richness across the Cerrado. High richness areas in northern Cerrado, predicted by our models, are still poorly sampled, and biological surveys are warranted in that region. The south‐western region of the Cerrado exhibits high species richness and is also undergoing high levels of deforestation. Therefore, maintenance of existing reserves, establishment of ecological corridors among reserves, and creation of new reserves are urgently needed to ensure conservation of species in these areas.