Persistence and extirpation in invaded landscapes: patch characteristics and connectivity determine effects of non-native predatory fish on native salamanders

Studies have demonstrated negative effects of non-native, predatory fishes on native amphibians, yet it is still unclear why some amphibian populations persist, while others are extirpated, following fish invasion. We examined this question by developing habitat-based occupancy models for the long-toed salamander (Ambystoma macrodactylum) and non-native fish using survey data from 1,749 water bodies across 470 catchments in the Northern Rocky Mountains, USA. We first modeled the habitat associations of salamanders at 468 fishless water bodies in 154 catchments where non-native fish were historically, and are currently, absent from the entire catchment. We then applied this habitat model to the complete data set to predict the probability of salamander occupancy in each water body, removing any effect of fish presence. Finally, we compared field-observed occurrences of salamanders and fish to modeled probability of salamander occupancy. Suitability models indicated that fish and salamanders had similar habitat preferences, possibly resulting in extirpations of salamander populations from entire catchments where suitable habitats were limiting. Salamanders coexisted with non-native fish in some catchments by using marginal quality, isolated (no inlet or outlet) habitats that remained fishless. They rarely coexisted with fish within individual water bodies and only where habitat quality was highest. Connectivity of water bodies via streams resulted in increased probability of fish invasion and consequently reduced probability of salamander occupancy. These results could be used to identify and prioritize catchments and water bodies where control measures would be most effective at restoring amphibian populations. Our approach could be useful as a framework for improved investigations into questions of persistence and extirpation of native species when non-native species have already become established.     

Local and Landscape Effects of Introduced Trout on Amphibians in Historically Fishless Watersheds

Introduced trout have often been implicated in the decline of high-mountain amphibian populations, but few studies have attempted to understand whether fish stocking also influences the distribution and abundance of amphibians throughout entire mountain basins, including the remaining fishless lakes. We examined this relationship using the relative abundance of long-toed salamanders (Ambystoma macrodactylum) and Columbia spotted frogs (Rana luteiventris) in fish-containing and fishless lentic sites in basins with varying levels of historic fish stocking. All lentic waters were surveyed for fish and amphibians in 11 high-elevation basins in the Frank Church-River of No Return Wilderness, Idaho, between 1994 and 1999. We found introduced trout (Oncorhynchus clarki, O. mykiss, O. m. aguabonita) in 43 of the 101 sites, representing 90% of the total surface area of lentic water bodies available. At the scale of individual water bodies, after accounting for differences in habitat characteristics between fish-containing and fishless sites, the abundance of amphibians at all life stages was significantly lower in lakes with fish. At the basin scale, densities of overwintering life stages of amphibians were lower in the fishless sites of basins where more habitat was occupied by trout. Our results suggest that many of the remaining fishless habitats are too shallow to provide suitable breeding or overwintering sites for these amphibians and that current trout distributions may eventually result in the extirpation of amphibian populations from entire landscapes, including sites that remain in a fishless condition. Received 28 March 2000;Accepted 2 January 2001.     

Sub-alpine amphibian distributions related to species palatability to non-native salmonids in the Klamath mountains of northern California

The goal of this study was to examine how introduced trout influence the distributions and abundances of a sub‐alpine amphibian assemblage whose members display a variety of different life‐history and defence strategies. Our study was conducted in the sub‐alpine lentic habitats of three wilderness areas that form the core of the Klamath‐Siskiyou Bioregion of northern California, a biodiversity ‘hotspot’ that supports the highest diversity of sub‐alpine, lentic‐breeding amphibians in the western USA. These wilderness areas contain no native fishes, but all have been populated with non‐native trout for recreational fishing. Five of the eight amphibian species that occur in this region were sufficiently common to use in our study; these included one that breeds in both temporary and permanent waters and is palatable to fish (Pacific treefrog, Pseudacris regilla), two that breed primarily in permanent waters and are unpalatable to fish (western toad, Bufo boreas, and rough‐skinned newt, Taricha granulosa), and two that breed primarily in permanent waters and are palatable to fish (Cascades frog, Rana cascadae, and long‐toed salamander, Ambystoma macrodactylum). Based on life histories and predator defence strategies (i.e. palatable or not), we predicted that the three palatable species would likely be negatively correlated with introduced trout, but with P. regilla less impacted because of its use of both temporary and permanent waters. We predicted that B. boreas and T. granulosa would not be significantly correlated with introduced trout due to the lack of any predator/prey interactions between them. We surveyed 728 pond, lake, or wet meadow sites during the summers of 1999–2002, using timed gill‐net sets to measure trout occurrence and relative density, and visual encounter surveys to determine amphibian presence and abundance. We used semiparametric logistic regression models to quantify the effect of trout presence/absence and density on the probability of finding amphibian species in a water body while accounting for variation within and among the various lentic habitats sampled. The distributions of P. regilla, A. macrodactylum and R. cascadae were strongly negatively correlated with trout presence across all three wilderness areas. Ambystoma macrodactylum was 44 times more likely to be found in lakes without fish than in lakes with fish. Rana cascadae and P. regilla were 3.7 and 3.0 times more likely, respectively, to be found in fishless than fish‐containing waters. In contrast, the two unpalatable species were either uncorrelated (T. granulosa) or positively correlated (B. boreas) with fish presence. We found that the relative density of fish (catch per unit effort) was negatively correlated with the combined abundances of the three palatable amphibians, and also with both the length and the condition of the fish themselves. Our results are consistent with a compelling body of evidence that introduced fishes greatly alter the aquatic community structure of mountain lakes, ponds, and wet meadows.     

Persistence at distributional edges: Columbia spotted frog habitat in the arid Great Basin,USA

A common challenge in the conservation of broadly distributed, yet imperiled species is understanding which factors facilitate persistence at distributional edges, locations where populations are often vulnerable to extirpation due to changes in climate, land use, or distributions of other species. For Columbia spotted frogs (Rana luteiventris) in the Great Basin (USA), a genetically distinct population segment of conservation concern, we approached this problem by examining (1) landscape‐scale habitat availability and distribution, (2) water body‐scale habitat associations, and (3) resource management‐identified threats to persistence. We found that areas with perennial aquatic habitat and suitable climate are extremely limited in the southern portion of the species’ range. Within these suitable areas, native and non‐native predators (trout and American bullfrogs [Lithobates catesbeianus]) are widespread and may further limit habitat availability in upper‐ and lower‐elevation areas, respectively. At the water body scale, spotted frog occupancy was associated with deeper sites containing abundant emergent vegetation and nontrout fish species. Streams with American beaver (Castor canadensis) frequently had these structural characteristics and were significantly more likely to be occupied than ponds, lakes, streams without beaver, or streams with inactive beaver ponds, highlighting the importance of active manipulation of stream environments by beaver. Native and non‐native trout reduced the likelihood of spotted frog occupancy, especially where emergent vegetation cover was sparse. Intensive livestock grazing, low aquatic connectivity, and ephemeral hydroperiods were also negatively associated with spotted frog occupancy. We conclude that persistence of this species at the arid end of its range has been largely facilitated by habitat stability (i.e., permanent hydroperiod), connectivity, predator‐free refugia, and a commensalistic interaction with an ecosystem engineer. Beaver‐induced changes to habitat quality, stability, and connectivity may increase spotted frog population resistance and resilience to seasonal drought, grazing, non‐native predators, and climate change, factors which threaten local or regional persistence.     

Predation Risk Avoidance by Terrestrial Amphibians: The Role of Prey Experience and Vulnerability to Native and Exotic Predators

We studied avoidance, by four amphibian prey species (Rana luteiventris, Ambystoma macrodactylum, Pseudacris regilla, Tarichia granulosa), of chemical cues associated with native garter snake (Thamnophis elegans) or exotic bullfrog (R. catesbeiana) predators. We predicted that avoidance of native predators would be most pronounced, and that prey species would differ in the intensity of their avoidance based on relative levels of vulnerability to predators in the wild. Adult R. luteiventris (presumably high vulnerability to predation) showed significant avoidance of chemical cues from both predators, A. macrodactylum (intermediate vulnerability to predation) avoided T. elegans only, while P. regilla (intermediate vulnerability to predation) and T. granulosa (low vulnerability to predation) showed no avoidance of either predator. We assessed if predator avoidance was innate and/or learned by testing responses of prey having disparate levels of prior exposure to predators. Wild‐caught (presumably predator‐exposed) post‐metamorphic juvenile R. luteiventris and P. regilla avoided T. elegans cues, while laboratory‐reared (predator‐naive) conspecifics did not; prior exposure to R. catesbeiana was not related to behavioural avoidance among adult or post‐metamorphic juvenile wild‐reared A. macrodactylum and P. regilla. These results imply that (i) some but not all species of amphibian prey avoid perceived risk from garter snake and bullfrog predators, (ii) the magnitude of this response probably differs according to prey vulnerability to predation in the wild, and (iii) avoidance tends to be largely learned rather than innate. Yet, the limited prevalence and intensity of amphibian responses to predation risk observed herein may be indicative of either a relatively weak predator–prey relationship and/or the limited importance of predator chemical cues in this particular system.     

Habitat Restoration as a Means of Controlling Non-Native Fish in a Mojave Desert Oasis

Non‐native fish generally cause native fish decline, and once non‐natives are established, control or elimination is usually problematic. Because non‐native fish colonization has been greatest in anthropogenically altered habitats, restoring habitat similar to predisturbance conditions may offer a viable means of non‐native fish control. In this investigation we identified habitats favoring native over non‐native fish in a Mojave Desert oasis (Ash Meadows) and used this information to restore one of its major warm water spring systems (Kings Pool Spring). Prior to restoration, native fishes predominated in warm water (25–32°C) stream and spring‐pool habitat, whereas non‐natives predominated in cool water (≤23°C) spring‐pool and marsh/slack water habitat. Native Amargosa pupfish (Cyprinodon nevadensis) and Ash Meadows speckled dace (Rhinichthys osculus nevadensis) inhabited significantly faster mean water column velocities (MWCV) and greater total depth (TD) than non‐native Sailfin molly (Poecilia latipinna) and Mosquitofish (Gambusia affinis) in warm water stream habitat, and Ash Meadows speckled dace inhabited significantly faster water than non‐natives in cool water stream habitat. Modification of the outflow of Kings Pool Spring from marsh to warm water stream, with MWCV, TD, and temperature favoring native fish, changed the fish composition from predominantly non‐native Sailfin molly and Mosquitofish to predominantly Ash Meadows pupfish. This result supports the hypothesis that restoring spring systems to a semblance of predisturbance conditions would promote recolonization of native fishes and deter non‐native fish invasion and proliferation.     

The Potential Role of Stream Habitat Restoration in Facilitating Salmonid Invasions: A Habitat‐Hydraulic Modeling Approach

Many successful invasions have taken place in systems where harmful disturbance has changed habitat conditions. However, less attention has been paid to the role of habitat restoration, which modifies habitats and thus also has the potential to facilitate invasions. We examined whether in‐stream habitat restorations have the potential to either facilitate or resist invasion by two widely introduced non‐native stream salmonids, Salvelinus fontinalis Mitchill and Oncorhynchus mykiss Walbaum, in Finland. A physical habitat simulation system was used to calculate whether the habitat area for the target species increased or decreased following the restorations. For comparison, we also reported results for four native stream fish species. The simulations showed that the restored streams provided the highest amount of usable habitat area for the native species, particularly for Salmo salar L. and Gottus gobio L. However, it was interesting to note that the restorations significantly increased habitat quality for the two non‐native species, especially at low flows. Nevertheless, the non‐native species had the lowest amount of usable habitat area overall. The modeling results indicated that not only habitat destruction but also habitat restoration could contribute to the spread of non‐native species. Fisheries and wildlife managers should be aware of the possibility, when restoring habitats in order to preserve native ecosystems, that non‐native species could manage to gain a foothold in restored habitats and use them as population sources for further spread. Knowing the widespread negative effect of non‐native species, this risk should not be underestimated.     

Predator Generalization Decreases the Effect of Introduced Predators in the San Marcos Salamander,Eurycea nana

The introduction of novel predators into an environment can have detrimental consequences on prey species, especially if these species lack the ability to recognize these predators. One such species that may be negatively affected by introduced predators is the federally threatened San Marcos salamander (Eurycea nana). Previous research found that predator‐naïve (captive‐hatched) salamanders showed decreased activity in response to the chemical cues of both a native fish predator (Micropterus salmoides) and an introduced fish predator (Lepomis auritus), but not to a non‐predatory fish (Gambusia geiseri). We tested the hypothesis that E. nana recognized the introduced Lepomis (and other non‐native Lepomis) because they share chemical cues with other native congeneric Lepomis predators in the San Marcos River. We examined the antipredator response of predator‐naïve E. nana to chemical cues from (1) a sympatric native sunfish (Lepomis cyanellus; Perciformes: Centrarchidae); (2) a sympatric introduced sunfish (L. auritus); (3) an allopatric sunfish (Lepomis gibbosus); (4) a sympatric non‐native, non‐centrarchid cichlid (Herichthys cyanoguttatum; Perciformes: Cichlidae); and (5) a blank water control to determine whether individuals make generalizations about novel predators within a genus and across a family. Exposure to chemical cues from all fish predator treatments caused a reduction in salamander activity (antipredator response). Additionally, there were no differences in the antipredator responses to each predatory fish treatment. The similar responses to all sunfish treatments indicate that E. nana shows predator generalization in response to novel predators that are similar to recognized predators. Additionally, the antipredator response to H. cyanoguttatum indicates that predator generalization can occur among perciform families.     

Strategies for the conservation and management of isolated salmonid populations: lessons from Japanese streams

1. Endangered native populations of stream salmonids in Japan face three major threats: (i) negative interactions with introduced hatchery‐reared fish, (ii) fragmentation of habitat by impassable dams and (iii) recreational angling. 2. To prevent imminent extinction of many local populations, we evaluated these threats and possible conservation actions for red‐spotted masu salmon (Oncorhynchus masou ishikawae) and white‐spotted charr (Salvelinus leucomaenis japonicus) in the Fuji River system in central Japan. 3. Red‐spotted masu salmon and white‐spotted charr occupied only 0.73 and 2.4% of suitable thermal habitats, respectively, with masu salmon typically occupying habitats closer to human population centres. 4. Population viability analysis resulted in a 100‐year probability of extinction of 78.1% for masu salmon and 48.1% for charr. However, extinction risk of both species was predicted to be <5% if the carrying capacity increased from 141 to 303 for masu salmon and from 94 to 125 for charr, by allowing fish passage at the lower end of the habitat, and if annual adult survival rate increased by 0.04. Adult survival rate was the principal factor associated with population persistence. 5. To conserve isolated populations of stream‐dwelling salmonids, we recommend (i) assessing the distribution of remnant native and non‐native fish populations, (ii) that fishing regulations are modified to improve adult survival and population persistence and (iii) that fragmented reaches be reconnected to adjacent habitat, for example by removing or modifying artificial barriers to increase the carrying capacity of the isolated populations. Reconnection of fragmented reaches should, however, be avoided if it results in non‐native fish invading isolated populations.     

Species‐specific responses to wetland mitigation among amphibians in the Greater Yellowstone Ecosystem

Habitat loss and degradation are leading causes of biodiversity declines, therefore assessing the capacity of created mitigation wetlands to replace habitat for wildlife has become a management priority. We used single season occupancy models to compare the occurrence of larvae of four species of pond‐breeding amphibians in wetlands created for mitigation, wetlands impacted by road construction, and unimpacted reference wetlands along a highway corridor in the Greater Yellowstone Ecosystem, United States. Created wetlands were shallow and had less aquatic vegetation and surface area than impacted and reference wetlands. Occupancy of barred tiger salamander (Ambystoma mavortium) and boreal chorus frog (Pseudacris maculata) larvae was similar across wetland types, whereas boreal toads (Anaxyrus boreas) occurred more often in created wetlands than reference and impacted wetlands. However, the majority of created wetlands (>80%) dried partially or completely before amphibian metamorphosis occurred in both years of our study, resulting in heavy mortality of larvae and, we suspect, little to no recruitment. Columbia spotted frogs (Rana luteiventris), which require emergent vegetation that is not common in newly created wetlands, occurred commonly in impacted and reference wetlands but were found in only one created wetland. Our results show that shallow created wetlands with little aquatic vegetation may be attractive breeding areas for some amphibians, but may result in high mortality and little recruitment if they fail to hold water for the entire larval period.     

Macroinvertebrates as indicators of fish absence in naturally fishless lakes

1. Little is known about native communities in naturally fishless lakes in eastern North America, a region where fish stocking has led to a decline in these habitats. 2. Our study objectives were to: (i) characterise and compare macroinvertebrate communities in fishless lakes found in two biophysical regions of Maine (U.S.A.): kettle lakes in the eastern lowlands and foothills and headwater lakes in the central and western mountains; (ii) identify unique attributes of fishless lake macroinvertebrate communities compared to lakes with fish and (iii) develop a method to efficiently identify fishless lakes when thorough fish surveys are not possible. 3. We quantified macroinvertebrate community structure in the two physiographic fishless lake types (n = 8 kettle lakes; n = 8 headwater lakes) with submerged light traps and sweep nets. We also compared fishless lake macroinvertebrate communities to those in fish‐containing lakes (n = 18) of similar size, location and maximum depth. We used non‐metric multidimensional scaling to assess differences in community structure and t‐tests for taxon‐specific comparisons between lakes. 4. Few differences in macroinvertebrate communities between the two physiographic fishless lake types were apparent. Fishless and fish‐containing lakes had numerous differences in macroinvertebrate community structure, abundance, taxonomic composition and species richness. Fish presence or absence was a stronger determinant of community structure in our study than differences in physical conditions relating to lake origin and physiography. 5. Communities in fishless lakes were more speciose and abundant than in fish‐containing lakes, especially taxa that are large, active and free‐swimming. Families differing in abundance and taxonomic composition included Notonectidae, Corixidae, Gyrinidae, Dytiscidae, Aeshnidae, Libellulidae and Chaoboridae. 6. We identified six taxa unique to fishless lakes that are robust indicators of fish absence: Graphoderus liberus, Hesperocorixa spp., Dineutus spp., Chaoborus americanus, Notonecta insulata and Callicorixa spp. These taxa are collected most effectively with submerged light traps. 7. Naturally fishless lakes warrant conservation, because they provide habitat for a unique suite of organisms that thrive in the absence of fish predation.     

Association of amphibians with attenuation of ultraviolet-b radiation in montane ponds

Ambient ultraviolet-b (UV-B) radiation (280-320 nm) has increased at north-temperate latitudes in the last two decades. UV-B can be detrimental to amphibians, and amphibians have shown declines in some areas during this same period. We documented the distribution of amphibians and salmonids in 42 remote, subalpine and alpine ponds in Olympic National Park, Washington, United States. We inferred relative exposure of amphibian habitats to UV-B by estimating the transmission of 305- and 320-nm radiation in pond water. We found breeding Ambystoma gracile, A. macrodactylum and Rana cascadae at 33%, 31%, and 45% of the study sites, respectively. Most R. cascadae bred in fishless shallow ponds with relatively low transmission of UV-B. The relationship with UV-B exposure remained marginally significant even after the presence of fish was included in the model. At 50 cm water depth, there was a 55% reduction in incident 305-nm radiation at sites where breeding populations of R. cascadae were detected compared to other sites. We did not detect associations between UV-B transmission and A. gracile or A. macrodactylum. Our field surveys do not provide evidence for decline of R. cascadae in Olympic National Park as has been documented in Northern California, but are consistent with the hypothesis that the spatial distribution of R. cascadae breeding sites is influenced by exposure to UV-B. Substrate or pond depth could also be related to the distribution of R. cascadae in Olympic National Park.     

Effects of Tamarix removal on the community dynamics of riparian birds in a semiarid grassland

Invasion of riparian habitats by non‐native plants is a global problem that requires an understanding of community‐level responses by native plants and animals. In the Great Plains, resource managers have initiated efforts to control the eastward incursion of Tamarix as a non‐native bottomland plant (Tamarix ramosissima) along the Cimarron River in southwestern Kansas, United States. To understand how native avifauna interact with non‐native plants, we studied the effects of Tamarix removal on riparian bird communities. We compared avian site occupancy of three foraging guilds, abundance of four nesting guilds, and assessed community dynamics with dynamic, multiseason occupancy models across three replicated treatments. Community parameters were estimated for Tamarix‐dominated sites (untreated), Tamarix‐removal sites (treated), and reference sites with native cottonwood sites (Populus deltoides). Estimates of initial occupancy (ψ₂₀₀₆) for the ground‐to‐shrub foraging guild tended to be highest at Tamarix‐dominated sites, while initial occupancy of the upper‐canopy foraging and mid‐canopy foraging guilds were highest in the treated and reference sites, respectively. Estimates of relative abundance for four nesting guilds indicated that the reference habitat supported the highest relative abundance of birds overall, although the untreated habitat had higher abundance of shrub‐nesters than treated or reference habitats. Riparian sites where invasive Tamarix is dominant in the Great Plains can provide nesting habitat for some native bird species, with avian abundance and diversity that are comparable to remnant riparian sites with native vegetation. Moreover, presence of some native vegetation in Tamarix‐dominated and Tamarix‐removal sites may increase abundance of riparian birds such as cavity‐nesters. Overall, our study demonstrates that Tamarix may substitute for native flora in providing nesting habitat for riparian birds at the eastern edge of its North American range.     

Higher temperatures enhance the effects of invasive sportfish on mountain zooplankton communities

1. Decades of introductions of exotic sportfish to mountain lakes around the world have impoverished them biologically, and this may be exacerbated by global warming. We assessed the current status of invasive salmonids and native zooplankton communities in 34 naturally fishless lakes along an elevational gradient, which served as an environmental proxy for the expected effects of climate change. 2. Our main goal was to explore how climate‐related variables influence the effects of stocked salmonids on the total biomass, species richness and taxonomic composition of zooplankton. We predicted that warmer conditions would dampen the negative predatory effects of exotic brook trout (Salvelinus fontinalis) on zooplankton communities because more temperate lakes contain a greater diversity of potentially tolerant species. 3. Instead, we discovered that the persistence of stocked brook trout in the warmer lakes significantly amplified total zooplankton biomass and species richness. In colder and deeper lakes, zooplankton were relatively unaffected by S. fontinalis, which however persisted better in alpine lakes than at lower elevations after stocking practices were halted over two decades ago. Warmer lake conditions and higher concentrations of dissolved organic carbon (DOC) were significant primary drivers of zooplankton species turnover, both favouring greater species diversity. 4. Our findings of an ecological surprise involving potential synergistic positive effects of climate warming and exotic trout on native zooplankton communities presents a conundrum for managers of certain national mountain parks. Present mandates to eradicate non‐native trout and return the mountain lakes to their naturally fishless state may conflict with efforts to conserve biodiversity under a rapidly changing climate.     

Effects of habitat size and isolation on species immigration–extinction dynamics and community nestedness in a desert river system

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Invasive trout increase the climatic sensitivity of zooplankton communities in naturally fishless lakes

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Structural differentiation of the meiotic and mitotic chromosomes of the salamander,Ambystoma macrodactylum

The lampbrush chromosomes of the long-toed salamander, Ambystoma macrodactylum Baird, have been analysed and a map of the oocyte genome prepared. The location of C-bands and cold-induced-constrictions has been established in mitotic chromosomes and compared with the location of marker structures and chiasmata in several lampbrush bivalents. In the lampbrush chromosomes, C-bands are tentatively correlated with sphere-organizing loci and with regions of low chiasma frequency; cold-induced-constrictions are tentatively correlated with regions of high chiasma frequency. In general, in this salamander, C-bands do not coincide in position with cold-induced-constrictions. We have compared our results with those obtained by Callan (1966) in his investigation of chromosomes of the axolotl, Ambystoma mexicanum, and we present an analysis of the similarities and differences that are visible in the chromosome sets of these two ambystomatid species.     

The importance (or lack thereof) of niche divergence to the maintenance of a northern species complex: the case of the long‐toed salamander (Ambystoma macrodactylum Baird)

The relative importance of ecological vs. nonecological factors for the origin and maintenance of species is an open question in evolutionary biology. Young lineages – such as the distinct genetic groups that make up the ranges of many northern species – represent an opportunity to study the importance of ecological divergence during the early stages of diversification. Yet, few studies have examined the extent of niche divergence between lineages in previously glaciated regions and the role of ecology in maintaining the contact zones between them. In this study, we used tests of niche overlap in combination with ecological niche models to explore the extent of niche divergence between lineages of the long‐toed salamander (Ambystoma macrodactylum Baird) species complex and to determine whether contact zones correspond to (divergent) niche limits. We found limited evidence for niche divergence between the different long‐toed salamander lineages, substantial overlap in the predicted distribution of suitable climatic space for all lineages and range limits that are independent of niche limits. These results raise questions as to the importance of ecological divergence to the development of this widespread species complex and highlight the potential for non‐ecological factors to play a more important role in the maintenance of northern taxa.     

Temperature,invaders and patchy habitat interact to limit the distribution of a vulnerable freshwater fish

Interacting global‐change drivers such as invasive species and climate warming are likely to have major and potentially unexpected influences on aquatic ecosystems. In river networks, modified water temperature combined with patchy physical conditions will likely cause shifts in the amount and distribution of suitable habitat, with influential invasive species further altering habitat availability. We examined how distributions of a thermally sensitive galaxiid fish native to the alpine rivers of New Zealand, Galaxias paucispondylus, were influenced by these drivers using spatially extensive presence–absence electrofishing surveys of 46 sites spread over four subcatchments. A unimodal response to water temperature and an interaction with substratum size meant G. paucispondylus were limited to streams with average summer water temperatures between 10.6 and 13.8 °C and were absent when average substratum sizes were <36 mm, regardless of temperature. In addition, non‐native trout >150 mm long excluded G. paucispondylus, but were only found in streams with average summer water temperatures <10.6 °C. These influences of trout likely strengthened the unimodal temperature response of G. paucispondylus and led to a very small G. paucispondylus realized niche. When predicted temperature increases were applied to catchment models, G. paucispondylus distributions were patchy and variable across subcatchments. Moreover, local physical characteristics of river networks were particularly important because of the non‐linear and interactive influences of temperature and substratum size on the outcome of species interactions. Therefore, substratum sizes, water temperature and a non‐native predator combined to influence the distribution of this thermally sensitive fish, illustrating how the effects of climate warming will likely be strongly context‐dependent and interactive.     

Tandem repeat markers for population genetic studies of the protected California tiger salamander (<Emphasis Type="Italic">Ambystoma californiense</Emphasis>)

Habitat loss is the single greatest threat to persistence of the critically threatened California tiger salamander (Ambystoma californiense). To aid management plans that designate critical habitat for this species, I developed and characterized 21 tetranucleotide microsatellite markers using two native populations in Santa Barbara and Alameda Counties. Allelic variation and average heterozygosities were lower in the endangered Santa Barbara population (allele range 1–4, mean 2.4; H _O = 0.308 H _E = 0.288) compared with the threatened Alameda population (allele range 2–10, mean 6.7; H _O = 0.712, H _E = 0.722). In-depth population studies using these markers will provide vital information for plans to assign critical habitat that optimize gene flow among breeding populations, as well as for identifying non-native hybrid genotypes that threaten native A. californiense stocks. Beyond the conservation goals for A. californiense, the close phylogenetic relationships within the tiger salamander complex also suggest a broad utility for population studies using these markers.