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Temporally separated species are often thought to have limited competition over a shared resource. However, early arriving species may consume a limited resource such that later-arriving species have access to fewer resources and thus experience competitive effects, even if they are temporally separated (i.e., they experience legacy effects from the early species). The presence of a predator might affect potential legacy effects by influencing the behavior or survivorship of the early species. Using a mesocosm experiment, I examined whether the presence of nonnative Western Mosquitofish (Gambusia affinis) mediated legacy effects in the interaction of two temporally separated species of tadpoles, early arriving American Toads (Anaxyrus americanus) and late-arriving Bullfrogs (Rana catesbeiana). Anaxyrus americanus tadpoles reduced R. catesbeiana tadpole growth despite all A. americanus tadpoles metamorphosing 8 days before the introduction of R. catesbeiana tadpoles into the mesocosms (i.e., legacy effects). Gambusia affinis had limited effects on A. americanus (1 day delay in metamorphosis but no effect on survivorship or size at metamorphosis) and positive effects on R. catesbeiana (increased growth). There were no significant interactions between the A. americanus tadpole density and G. affinis treatments. In conclusion, I found evidence of significant legacy effects of A. americanus tadpoles on R. catesbeiana tadpoles, but no evidence that G. affinis mediated the legacy effects.

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Forests are major carbon (C) sinks, but their ability to sequester C and thus mitigate climate change, varies with the environment, disturbance regime, and biotic interactions. Herbivory by invasive, nonnative ungulates can have profound ecosystem effects, yet its consequences for forest C stocks remain poorly understood. We determined the impact of invasive ungulates on C pools, both above- and belowground (to 30 cm), and on forest structure and diversity using 26 paired long-term (>20 years) ungulate exclosures and adjacent unfenced control plots located in native temperate rainforests across New Zealand, spanning 36-41° S. Total ecosystem C was similar between ungulate exclosure (299.93 ± 25.94 Mg C ha-1 ) and unfenced control (324.60 ± 38.39 Mg C ha-1 ) plots. Most (60%) variation in total ecosystem C was explained by the biomass of the largest tree (mean diameter at breast height [dbh]: 88 cm) within each plot. Ungulate exclusion increased the abundance and diversity of saplings and small trees (dbh ≥2.5, <10 cm) compared with unfenced controls, but these accounted for ~5% of total ecosystem C, demonstrating that a few, large trees dominate the total forest ecosystem C but are unaffected by invasive ungulates at a timescale of 20-50 years. However, changes in understory C pools, species composition, and functional diversity did occur following long-term ungulate exclusion. Our findings suggest that, although the removal of invasive herbivores may not affect total forest C at the decadal scale, major shifts in the diversity and composition of regenerating species will have longer term consequences for ecosystem processes and forest C.

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Isolated, seasonal wetlands within agricultural landscapes are important ecosystems. However, they are currently experiencing direct and indirect effects of agricultural management surrounding them. Because wetlands provide important ecosystem services, it is crucial to determine how these factors affect ecological communities. Here, we studied the long-term effects of land-use intensification, cattle grazing, prescribed fires, and their interactions on wetland plant diversity, community dynamics, and functional diversity. To do this, we used vegetation and trait data from a 14-year-old experiment on 40 seasonal wetlands located within seminatural and intensively managed pastures in Florida. These wetlands were allocated different grazing and prescribed fire treatments (grazed vs. ungrazed, burned vs. unburned). Our results showed that wetlands within intensively managed pastures have lower native plant diversity, floristic quality, evenness, and higher nonnative species diversity and exhibited the most resource-acquisitive traits. Wetlands embedded in intensively managed pastures were also characterized by lower species turnover over time. We found that 14 years of cattle exclusion reduced species diversity in both pasture management intensities and had no effect on floristic quality. Fenced wetlands exhibited lower functional diversity and experienced a higher rate of community change, both due to an increase in tall, clonal, and palatable grasses. The effects of prescribed fires were often dependent on grazing treatment. For instance, prescribed fires increased functional diversity in fenced wetlands but not in grazed wetlands. Our study suggests that cattle exclusion and prescribed fires are not enough to restore wetlands in intensively managed pastures and further highlights the importance of not converting seminatural pastures to intensively managed pastures. Our study also suggests that grazing levels applied in seminatural pastures maintained high plant diversity and prevented tree and shrub encroachment and that in the absence of grazing, prescribed fire became crucial to maintaining higher species evenness.

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The movement of plant species across the globe exposes native communities to new species introductions. While introductions are pervasive, two aspects of variability underlie patterns and processes of biological invasions at macroecological scales. First, only a portion of introduced species become invaders capable of substantially impacting ecosystems. Second, species that do become invasive at one location may not be invasive in others; impacts depend on invader abundance and recipient species and conditions. Accounting for these phenomena is essential to accurately understand the patterns of plant invasion and explain the idiosyncratic results reflected in the literature on biological invasions. The lack of community-level richness and the abundance of data spanning broad scales and environmental conditions have until now hindered our understanding of invasions at a macroecological scale. To address this limitation, we leveraged quantitative surveys of plant communities in the USA and integrated and harmonized nine datasets into the Standardized Plant Community with Introduced Status (SPCIS) database. The database contains 14,056 unique taxa identified within 83,391 sampling units, of which 52.6% have at least one introduced species. The SPCIS database includes comparable information on plant species occurrence, abundance, and native status across the 50 U.S. States and Puerto Rico. SPCIS can be used to answer macro-scale questions about native plant communities and interactions with invasive plants. There are no copyright restrictions on the data, and we ask the users of this dataset to cite this paper, the respective paper(s) corresponding to the dataset sampling design (all references are provided in Data S1: Metadata S1: Class II-B-2), and the references described in Data S1: Metadata S1: Class III-B-4 as applicable to the dataset being utilized.

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Relatively few studies have attempted to resolve the pathways through which the effects of biodiversity on ecosystem functioning cascade from one trophic level to another. Here, we manipulated the richness of habitat-forming seaweeds in a western Atlantic estuary to explore how changes in foundation species diversity affect the structure and functioning of the benthic consumer communities that they support. Structural equation modeling revealed that macroalgal richness enhanced invertebrate abundance, biomass, and diversity, both directly by changing the quality and palatability of the foundational substrate and indirectly by increasing the total biomass of available habitat. Consumer responses were largely driven by a single foundational seaweed, although stronger complementarity among macroalgae was observed for invertebrate richness. These findings with diverse foundational phyla extend earlier inferences from terrestrial grasslands by showing that biodiversity effects can simultaneously propagate through multiple independent pathways to maintain animal foodwebs. Our work also highlights the potential ramifications of human-induced changes in marine ecosystems.

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Knowledge of a species' phenology can assist with timing accurate surveys to detect presence and density in a novel environment. Trichoferus campestris (Faldermann) (Coleoptera: Cerambycidae) has been found in the United States since the 1990s, but its biology and behavior remain poorly understood. This study investigated the phenology of T. campestris in Minneapolis and St. Paul, Minnesota, USA, based on local temperature data. In the summers of 2019 and 2020, 30 and 40 pheromone-baited traps, respectively, were deployed in trees in suburban parks to monitor capture of local individuals. Traps were suspended from branches of mature trees selected at random, with nine genera of trees selected over both years. Early, peak, and late adult abundance were characterized, and the impact of tree genus in which each trap was hung was evaluated. Abundance was found to be unimodal both summers with a peak around 650 degree days (base 10°C) in early July. Significantly more adult T. campestris were caught in traps hung in trees of the genus Tilia than in trees of the genus Quercus. These findings are important first steps to improving monitoring of T. campestris presence and conducting risk assessments.

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SignificanceThe introduction of trees outside their native ranges has greatly expanded the potential ranges of their pathogens and insect pests, which risk spilling over and impacting native flora. However, we often lack a strong understanding of the host, climatic, and geographic factors that allow pests to establish outside their hosts' native ranges. Using global datasets of pest occurrences and the native and nonnative ranges of tree hosts, we show there are strong generalizable trends controlling pest occurrences and can predict the occurrence of pests outside their hosts' native ranges with >75% accuracy. Our modeling framework offers a powerful tool to identify future invasive pest species and the ecological mechanisms controlling the accumulation of pests outside their hosts' native ranges.

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Originally endemic to South America, the nine-banded armadillo (Dasypus novemcinctus) has recently expanded its range northward to Illinois. With this range expansion comes concern regarding potential incoming pathogens; our research, conducted during 2012-2020, consisted of screening armadillos for the presence of helminths, Trypanosoma cruzi, and Mycobacterium leprae. We screened for the presence of T. cruzi and M. leprae, 2 pathogens known to infect humans, using polymerase chain reaction and enzyme-linked immunosorbent assay, respectively. All 80 samples screened for T. cruzi and all 25 samples screened for M. leprae were negative. No parasite specific to the nine-banded armadillo, such as Aspidodera sogandaresi, was detected. This lack of infection is consistent with the idea that animals may be isolated from their common parasites during periods of range expansion. Lack of infection by T. cruzi in an endemic area suggests that these mammals may not be exposed to the infective stages at this early phase of their colonization. Presently, the armadillo has become established in Illinois, yet they have not introduced their parasites into the area. Our study represents the first effort to document the parasitological record of the expanding armadillo within 30 yr of their initial record in Illinois and the American Midwest. This helminthological record of armadillos in Illinois sets the timeline to observe the establishment of A. sogandaresi in the Midwest.

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Large-scale movement of organisms across their habitable range, or migration, is an important evolutionary process that can shape genetic diversity and influence the adaptive spread of alleles. Although human migrations have been studied in great detail with modern and ancient genomes, recent anthropogenic influence on reducing the biogeographical constraints on the migration of nonnative species has presented opportunities in several study systems to ask the questions about how repeated introductions shape genetic diversity in the introduced range. We present an extensive overview of population structure of North American Arabidopsis thaliana by studying a set of 500 whole-genome sequenced and over 2,800 RAD-seq genotyped individuals in the context of global diversity represented by Afro-Eurasian genomes. We use methods based on haplotype and rare-allele sharing as well as phylogenetic modeling to identify likely sources of introductions of extant N. American A. thaliana from the native range in Africa and Eurasia. We find evidence of admixture among the introduced lineages having increased haplotype diversity and reduced mutational load. We also detect signals of selection in immune-system-related genes that may impart qualitative disease resistance to pathogens of bacterial and oomycete origin. We conclude that multiple introductions to a nonnative range can rapidly enhance the adaptive potential of a colonizing species by increasing haplotypic diversity through admixture. Our results lay the foundation for further investigations into the functional significance of admixture.

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Soil seed banks represent reservoirs of diversity in the soil that may increase resilience of communities to global changes. Two global change factors that can dramatically alter the composition and diversity of aboveground communities are nutrient enrichment and increased rainfall. In a full-factorial nutrient and rainfall addition experiment in an annual Californian grassland, we asked whether shifts in aboveground composition and diversity were reflected in belowground seed banks. Nutrient and rainfall additions increased exotic and decreased native abundances, while rainfall addition increased exotic richness, both in aboveground communities and seed banks. Under nutrient addition, forbs and short-statured plants were replaced by grasses and tall-statured species, both above and below ground, and whole-community responses to the treatments were similar. Structural equation models indicated that especially nutrient addition effects on seed banks were largely indirect via aboveground communities. However, rainfall addition also had a direct negative effect on native species richness and abundance of species with high specific leaf area (SLA) in seed banks, showing that seed banks are sensitive to the direct effects of temporary increases in rainfall. Our findings highlight the vulnerability of seed banks in annual, resource-poor grasslands to shifts in compositional and trait changes in aboveground communities and show how invasion of exotics and depletion of natives are critical for these above-belowground compositional shifts. Our findings suggest that seed banks have limited potential to buffer resource-poor annual grasslands from the community changes caused by resource enrichment.

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Invasive plant species (IAS), with their numerous negative ecological, health, and economic impacts, represent one of the greatest conservation challenges in the world. Reducing the negative impacts and potentially exploiting the biomass of these plant species can significantly contribute to sustainable management, protect biodiversity, and create a healthy environment. Therefore, the main objective of this study was to evaluate the nutritional potential, phytochemical status, and antioxidant capacity of nine alien invasive plant species: Abutilon theophrasti, Amaranthus retroflexus, Ambrosia artemisiifolia, Datura stramonium, Erigeron annuus, Galinsoga ciliata, Reynoutria japonica, Solidago gigantea, and Sorghum halepense. Multivariate statistical methods such as cluster and PCA were performed to determine possible connections and correlations among selected IAS depending on the phytochemical content. According to the obtained results, R. japonica was notable with the highest content of vitamin C (38.46 mg/100 g FW); while E. annuus (1365.92 mg GAE/100 g FW) showed the highest values of total polyphenolic compounds. A. retroflexus was characterized by the highest content of total chlorophylls (0.26 mg/g) and antioxidant capacity (2221.97 µmol TE/kg). Therefore, it can be concluded that the selected IAS represent nutrient-rich plant material with significant potential for the recovering of bioactive compounds.

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Understanding the mechanisms of spatial variation of biological invasions, across local-to-global scales, has been a major challenge. The importance of evolutionary history for invasion dynamics was noted by Darwin, and several studies have since considered how biodiversity of source and recipient regions can influence the probability of invasions. For over a century, the Panama Canal has connected water bodies and biotas with different evolutionary histories, and created a global shipping hot spot, providing unique opportunities to test mechanisms that affect invasion patterns. Here, we test for asymmetry in both the extent of invasions and predation effects, a possible mechanism of biotic resistance, between two tropical oceans at similar latitudes. We estimated nonnative species (NNS) richness for sessile marine invertebrates, using standardized field surveys and literature synthesis, to examine whether invasions are asymmetrical, with more NNS present in the less diverse Pacific compared to the Atlantic. We also experimentally tested whether predation differentially limits the abundance and distribution of these invertebrates between oceans. In standardized surveys, observed total NNS richness was higher in the Pacific (18 NNS, 30% of all Pacific species) than the Atlantic (11 NNS, 13% of all Atlantic species). Similarly, literature-based records also display this asymmetry between coasts. When considering only the reciprocal exchange of NNS between Atlantic and Pacific biotas, NNS exchange from Atlantic to Pacific was eightfold higher than the opposite direction, exceeding the asymmetry predicted by random exchange based simply on differences of overall diversity per region. Predation substantially reduced biomass and changed NNS composition in the Pacific, but no such effects were detected on the Atlantic coast. Specifically, some dominant NNS were particularly susceptible to predation in the Pacific, supporting the hypothesis that predation may reduce the abundance of certain NNS here. These results are consistent with predictions that high diversity in source regions, and species interactions in recipient regions, shape marine invasion patterns. Our comparisons and experiments across two tropical ocean basins, suggest that global invasion dynamics are likely driven by both ecological and evolutionary factors that shape susceptibility to and directionality of invasions across biogeographic scales.

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Biotic resistance is often posited, but rarely known, to be the cause of invasion failure. Competition and predation are the most frequently identified processes that may prevent or limit the establishment of nonnative species. Interactions between nonnative and native species that involve intraguild predation (IGP) are very common in nature, although theory predicts most IGP systems should be unstable and lead to extinction. If this prediction is true, the frequency of invasion failures due to IGP may be underappreciated because of their fleeting nature and, thus, studies of unstable IGP systems are lacking, despite the opportunities they offer for understanding the factors affecting their unstable dynamics. We investigated a failed invasion involving an IGP relationship. In Florida, the guppy (Poecilia reticulata), a worldwide invader, fails to establish in the presence of eastern mosquitofish (Gambusia holbrooki). We tested whether and how resident mosquitofish cause guppy invasion failure using replicated mesocosm and aquarium trials. Both the predator and competitor components of the IGP relationship were strongly asymmetrical, with large impacts on guppies. We identified two effects, direct consumption of neonates and aggressive interference competition, that limited survival and recruitment. The highly unstable nature of this IGP relationship is the primary cause of the failure of the guppy to establish in Florida. Our study shows that the transient nature of an ephemeral IGP relationship can yield important insights into the underlying causes of invasion failure, including the role of strong biotic resistance.

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Philophthalmus is a genus of globally distributed parasitic eye flukes with some members of the genus found in disparate locales. In particular, Philophthalmus gralli, a zoonotic trematode, appears to be a relatively new introduction to the Americas, facilitated by spillover from the invasive snails Melanoides tuberculata (red-rimmed melania) and Tarebia granifera (quilted melania), which were introduced via the aquarium trade, and perhaps furthered by avian dispersal. Given that two known intermediate hosts of Philophthalmus flukes are actively expanding their range as a result of human activities, we hypothesize that this spread is also associated with the spread of Philophthalmus flukes. To address this, we systematically reviewed the literature and examined whether the global expansion of P. gralli flukes is associated with the spread of invasive snails M. tuberculata and T. granifera. Here, we show that (1) specimens of P. gralli are only found in intermediate snail hosts M. tuberculata or T. granifera, suggesting intermediate host specificity for these 2 species, and (2) specimens of P. gralli have rarely been found outside the ranges (native and introduced) of M. tuberculata or T. granifera. Given the importance of distribution information of parasites in the role of identifying parasite invasions, we also review the known distribution of all Philophthalmus species. Considering recent outbreaks in humans and wild and domestic animal species, the continued spread of Philophthalmus presents a potential threat to veterinary and public health and conservation.

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Climate change affects ecological processes and interactions, including parasitism. Because parasites are natural components of ecological systems, as well as agents of outbreak and disease-induced mortality, it is important to summarize current knowledge of the sensitivity of parasites to climate and identify how to better predict their responses to it. This need is particularly great in marine systems, where the responses of parasites to climate variables are less well studied than those in other biomes. As examples of climate's influence on parasitism increase, they enable generalizations of expected responses as well as insight into useful study approaches, such as thermal performance curves that compare the vital rates of hosts and parasites when exposed to several temperatures across a gradient. For parasites not killed by rising temperatures, some simple physiological rules, including the tendency of temperature to increase the metabolism of ectotherms and increase oxygen stress on hosts, suggest that parasites' intensity and pathologies might increase. In addition to temperature, climate-induced changes in dissolved oxygen, ocean acidity, salinity, and host and parasite distributions also affect parasitism and disease, but these factors are much less studied. Finally, because parasites are constituents of ecological communities, we must consider indirect and secondary effects stemming from climate-induced changes in host-parasite interactions, which may not be evident if these interactions are studied in isolation.

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Managing the world's freshwater supply to meet societal and environmental needs in a changing climate is one of the biggest challenges for the 21st century. Dams provide water security; however, the allocation of dwindling water supply among reservoirs could exacerbate or ameliorate the effects of climate change on aquatic communities. Here, we show that the relative sensitivity of river thermal regimes to direct impacts of climate change and societal decisions concerning water storage vary substantially throughout a river basin. In the absence of interspecific interactions, future Colorado River temperatures would appear to benefit both endemic and nonnative fish species. However, endemic species are already declining or extirpated in locations where their ranges overlap with warmwater nonnatives and changes in water storage may lead to warming in some of the coolest portions of the river basin, facilitating further nonnative expansion. Integrating environmental considerations into ongoing water storage negotiations may lead to better resource outcomes than mitigating nonnative species impacts after the fact.

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The addition of nonnative species and loss of native species has modified the composition of communities globally. Although changes in ß-diversity have been well documented, there is a need for studies incorporating multiple time periods, more than one dimension of biodiversity, and inclusion of nestedness and turnover components to understand the underlying mechanisms structuring community composition and assembly. Here, we examined temporal changes in functional dissimilarity of fish communities of the Laurentian Great Lakes and compared these changes to those of taxonomic dissimilarity by decade from 1870 to 2010. Jaccard-derived functional dissimilarity index was used to quantify changes in functional ß-diversity within communities, between all possible pairs of communities, and using a multiple-site index among all communities. ß-diversity was partitioned into components of nestedness and turnover, and changes were examined over time. Similar to patterns in taxonomic dissimilarity, each community functionally differentiated from the historical community of 1870, with Lake Superior changing the most (~24%) and Lake Ontario the least (~14%). Although communities have become taxonomically homogenized, functional ß-diversity among all communities has increased over time, indicating functional differentiation. This is likely due to functional similarity between the communities being historically high (i.e., ~88% similar in 1870). The higher taxonomic relative to functional turnover indicates that the species being replaced between communities are functionally redundant, which could occur given the harsh environmental conditions of the region and/or as a result of the recent glacial history of the region. High functional nestedness across communities reflects dispersal limitations, with smaller communities being functional subsets of large communities closer to source populations. The functional differentiation observed is likely due to nonnative species with functional traits unique to the region establishing or the loss of functionally redundant native species; however, it is important to note that patterns of homogenization were periodically observed through time. Our study demonstrates the possible factors regulating diversity in the Laurentian Great Lakes fish communities, that patterns of taxonomic and functional ß-diversity are dynamic over time and vary in the magnitude and direction of change, and that taxonomic diversity should not be used to predict changes in functional diversity.

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The introduction of nonnative species is one of the most critical problems facing freshwater systems today. The rivers of the Great Basin (USA) have been particularly imperilled by nonnative species introductions and represent a valuable location to study the dietary trends of native and nonnative fishes in isolated, endorheic systems. We collected fish from 23 sites, spanning three Great Basin watersheds (Carson, Humboldt and Bear Rivers) and two elevation categories (upland and lowland). Only a single species (speckled dace Rhinichthys osculus) occurred in both elevation zones. Diet item analyses of over 500 fish stomachs indicated significant dietary overlaps between native and nonnative fishes and detailed dietary selectivity for all species. This finding, along with the low species diversity observed in the region, suggests low dietary niche diversity, which could have the potential to amplify the competitive impacts of nonnatives on native species. In upland sites, nonnative trouts were the dominant invaders, while in lowland sites warm-water nonnatives were prevalent. The management implications we recommend based on our results urge for continued monitoring of water temperature and species occurrences to predict if dietary overlaps observed in this study are likely to change in the future. SIGNIFICANCE STATEMENT: The Great Basin is an ideal endorheic region to study dietary trends in native and nonnative fishes. These trends are important in predicting competitive interactions among fishes. By looking at the diets of fishes within this region we were able to identify multiple significant overlaps among native and nonnative fishes. These results represent a baseline for future studies in the region as well as being comparable to other regions with similar invasive/native overlaps.

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Coastal habitats in Chile are hypothesized to support a number of diadromous fish species. The objective of this study was to document migratory life histories of native galaxiids and introduced salmonids from a wide latitudinal range in Chilean Patagonia (39-48°S). Otolith microchemistry data were analysed using a recursive partitioning approach to test for diadromy. Based on annular analysis of Sr:Ca ratios, a diadromous life history was suggested for populations of native Aplochiton taeniatus, A. marinus, and Galaxias maculatus. Lifetime residency in freshwater was suggested for populations of A. zebra and G. platei. Among introduced salmonids, populations of Oncorhynchus tshawytscha and O. kisutch exhibited patterns consistent with anadromy, whereas the screened population of O. mykiss appeared restricted to freshwater. Salmo trutta exhibited variable patterns suggesting freshwater residency and possibly anadromy in one case. The capacity and geographic scope of hydropower development is increasing and may disrupt migratory routes of diadromous fishes. Identification of diadromous species is a critical first step for preventing their loss due to hydropower development.

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Temperature has a substantial effect on both the physiology and behavior of ectothermic animals such as lizards. Physiology and behavior can also be influenced by ontogenetic and sex differences, but these effects are largely understudied in lizards. We examined ontogenetic and sex-based differences in thermal tolerances, preferred temperature, and temperature-dependent evaporative water loss rates in Italian Wall Lizards, Podarcis siculus, collected from an introduced population near Los Angeles, California, USA that were acclimated to laboratory conditions. Podarcis siculus has been introduced to multiple localities in the USA and the Mediterranean region and has demonstrated remarkable ability to adapt to novel climatic conditions. In the California population, adults of both sexes had a higher critical thermal maximum (CTmax) than juveniles, and adult females had a lower critical thermal minimum (CTmin) than juveniles and adult males. Thus, adult females had a significantly wider thermal breadth (CTmax - CTmin) compared to adult males and juveniles. Mass-specific evaporative water loss was higher in juveniles compared to adult males at intermediate temperatures. There was no significant difference among groups for preferred temperature. This implies that thermal tolerance, a physiological characteristic, varies with age and sex for this population, whereas thermal preference, a behavioral characteristic, does not. Interestingly, CTmin for all age and sex classes was above temperatures likely experienced by some nonnative populations in winter, suggesting individuals need to find urban thermal retreats. These results add to the growing literature demonstrating that thermal tolerances and breadths can vary between sexes and across age classes in squamate species.

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