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Accumulating evidence in ecology: Once is not enough

Many published studies in ecological science are viewed as stand-alone investigations that purport to provide new insights into how ecological systems behave based on single analyses. But it is rare for results of single studies to provide definitive results, as evidenced in current discussions of the “reproducibility crisis” in science. The key step in science is the comparison of hypothesis-based predictions with observations, where the predictions are typically generated by hypothesis-specific models. Repeating this step allows us to gain confidence in the predictive ability of a model, and its corresponding hypothesis, and thus to accumulate evidence and eventually knowledge. This accumulation may occur via an ad hoc approach, via meta-analyses, or via a more systematic approach based on the anticipated evolution of an information state. We argue the merits of this latter approach, provide an example, and discuss implications for designing sequences of studies focused on a particular question. We conclude by discussing current data collection programs that are pre-adapted to use this approach and argue that expanded use would increase the rate of learning in ecology, as well as our confidence in what is learned.

Ecology and Evolution↗

Deconstructing cardiovascular and coagulation-related traits links dietary ecology to multi-functional snake venom specificity

Animal venoms vary greatly in compositional complexity, where complex venoms are hypothesized to be maintained by greater dietary breadth. Beyond explaining venom composition, the dietary breadth hypothesis predicts that these more complex venoms should show greater functional breadth in terms of overall toxicity and by disrupting multiple prey physiological processes. We evaluate these predictions with six distinct physiological assays of cardiac and blood clotting functions and compared the effects of venoms from snake species with a range of taxonomic dietary diversity levels. We compared the taxonomic dietary generalists Agkistrodon piscivorus and Sistrurus miliarius to taxa with varying taxonomic specialization, namely Ag. contortrix and Crotalus adamanteus , and Azemiops feae . Comparing fish thrombocyte and mammal platelet aggregation and fibrin clot formation, only species with broader diets disrupted both fish and mammal hemostatic function. Venom of Ag. piscivorus , a uniquely fish-eating species, was the most disruptive of zebrafish heart rate, thrombocyte activation, vascular permeability, and clotting after injury. Sistrurus miliarius was also highly toxic, whereas mammal-specialists’ venoms scarcely altered zebrafish physiology. Our results support the hypothesis that dietary breadth selects functionally complex venoms. Understanding venom gene evolution, snakebite symptoms, and searching for therapeutics in venom should be guided by evolutionary ecology.

Evolution↗

Ecological divergence of wild birds drives avian influenza spillover and global spread

The diversity of influenza A viruses (IAV) is primarily hosted by two highly divergent avian orders: Anseriformes (ducks, swans and geese) and Charadriiformes (gulls, terns and shorebirds). Studies of IAV have historically focused on Anseriformes, specifically dabbling ducks, overlooking the diversity of hosts in nature, including gull and goose species that have successfully adapted to human habitats. This study sought to address this imbalance by characterizing spillover dynamics and global transmission patterns of IAV over 10 years at greater taxonomic resolution than previously considered. Furthermore, the circulation of viral subtypes in birds that are either host-adapted (low pathogenic H13, H16) or host-generalist (highly pathogenic avian influenza—HPAI H5) provided a unique opportunity to test and extend models of viral evolution. Using Bayesian phylodynamic modelling we uncovered a complex transmission network that relied on ecologically divergent bird hosts. The generalist subtype, HPAI H5 was driven largely by wild geese and swans that acted as a source for wild ducks, gulls, land birds, and domestic geese. Gulls were responsible for moving HPAI H5 more rapidly than any other host, a finding that may reflect their long-distance, pelagic movements and their immuno-naïve status against this subtype. Wild ducks, long viewed as primary hosts for spillover, occupied an optimal space for viral transmission, contributing to geographic expansion and rapid dispersal of HPAI H5. Evidence of inter-hemispheric dispersal via both the Pacific and Atlantic Rims was detected, supporting surveillance at high latitudes along continental margins to achieve early detection. Both neutral (geographic expansion) and non-neutral (antigenic selection) evolutionary processes were found to shape subtype evolution which manifested as unique geographic hotspots for each subtype at the global scale. This study reveals how a diversity of avian hosts contribute to viral spread and spillover with the potential to improve surveillance in an era of rapid global change.

PLoS Pathogens↗

Evolving wildlife management cultures of governance through Indigenous Knowledges and perspectives

Within governance agencies, academia, and communities alike, there are increasing calls to recognize the value and importance of culture within social-ecological systems and to better implement Indigenous sciences in research, policy, and management. Efforts thus far have raised questions about the best ethical practices to do so. Engaging with plural worldviews and perspectives on their own terms reflects cultural evolutionary processes driving paradigm shifts in 3 fundamental areas of natural resource management: conceptualizations of natural resources and ecosystems, processes of public participation and governance, and relationships with Indigenous Peoples and communities with differing worldviews. We broadly describe evolution toward these paradigm shifts in fish and wildlife management. We then use 3 case studies to illustrate the ongoing cultural evolution of relationships between wildlife management and Indigenous practices within specific historical and social-ecological contexts and reflect on common barriers to appropriately engaging with Indigenous paradigms and lifeways. Our case studies highlight 3 priorities that can assist the field of wildlife management in achieving the changes necessary to bridge incommensurable worldviews: acknowledging and reconciling historical legacies and their continued power dynamics as part of social-ecological systems, establishing governance arrangements that move beyond attempts to extract cultural information from communities to integrate Indigenous Knowledges into dominant management paradigms, and engaging in critical reflexivity and reciprocal, accountable relationship building. Implementing these changes will take time and a commitment to processes that may initially feel uncomfortable and unfamiliar but have potential to be transformative. Ethical and culturally appropriate methods to include plural and multivocal perspectives and worldviews on their own terms are needed to transform wildlife management to achieve more effective and just management outcomes for all.

Journal of Wildlife Management↗

Microbial aggregates within tissues infect a diversity of corals throughout the Indo-Pacific

Coral reefs are highly diverse ecosystems where symbioses play a pivotal role. Corals contain cell-associated microbial aggregates (CAMA), yet little is known about how widespread they are among coral species or the nature of the symbiotic relationship. Using histology, we found CAMA within 24 species of corals from 6 genera from Hawaii, American Samoa, Palmyra, Johnston Atoll, Guam, and Australia. Prevalence (%) of infection varied among coral genera: Acropora, Porites, and Pocillopora were commonly infected whereas Montipora were not. Acropora from the Western Pacific were significantly more likely to be infected with CAMA than those from the Central Pacific, whereas the reverse was true for Porites. Compared with apparently healthy colonies, tissues from diseased colonies were significantly more likely to have both surface and basal body walls infected. The close association of CAMA with host cells in numerous species of apparently healthy corals and lack of associated cell pathology reveals an intimate agent-host association. Furthermore, CAMA are Gram negative and in some corals may be related to chlamydia or rickettsia. We propose that CAMA in adult corals are facultative secondary symbionts that could play an important ecological role in some dominant coral genera in the Indo-Pacific. CAMA are important in the life histories of other animals, and more work is needed to understand their role in the distribution, evolution, physiology, and immunology of reef corals.

Marine Ecology Progress Series↗

Representing the acquisition and use of energy by individuals in agent-based models of animal populations

1. Agent-based models (ABMs) are widely used to predict how populations respond to changing environments. As the availability of food varies in space and time, individuals should have their own energy budgets, but there is no consensus as to how these should be modelled. Here, we use knowledge of physiological ecology to identify major issues confronting the modeller and to make recommendations about how energy budgets for use in ABMs should be constructed. 2. Our proposal is that modelled animals forage as necessary to supply their energy needs for maintenance, growth and reproduction. If there is sufficient energy intake, an animal allocates the energy obtained in the order: maintenance, growth, reproduction, energy storage, until its energy stores reach an optimal level. If there is a shortfall, the priorities for maintenance and growth/reproduction remain the same until reserves fall to a critical threshold below which all are allocated to maintenance. Rates of ingestion and allocation depend on body mass and temperature. We make suggestions for how each of these processes should be modelled mathematically. 3. Mortality rates vary with body mass and temperature according to known relationships, and these can be used to obtain estimates of background mortality rate. 4. If parameter values cannot be obtained directly, then values may provisionally be obtained by parameter borrowing, pattern-oriented modelling, artificial evolution or from allometric equations. 5. The development of ABMs incorporating individual energy budgets is essential for realistic modelling of populations affected by food availability. Such ABMs are already being used to guide conservation planning of nature reserves and shell fisheries, to assess environmental impacts of building proposals including wind farms and highways and to assess the effects on nontarget organisms of chemicals for the control of agricultural pests.

Methods in Ecology and Evolution↗

Conspecific reproductive success and breeding habitat selection: Implications for the study of coloniality

Habitat selection is a crucial process in the life cycle of animals because it can affect most components of fitness. It has been proposed that some animals cue on the reproductive success of conspecifics to select breeding habitats. We tested this hypothesis with demographic and behavioral data from a 17-yr study of the Black-legged Kittiwake ( Rissa tridactyla ), a cliff-nesting seabird. As the hypothesis assumes, the Black-legged Kittiwake nesting environment was patchy, and the relative quality of the different patches (i.e., breeding cliffs) varied in time. The average reproductive success of the breeders of a given cliff was predictable from one year to the next, but this predictability faded after several years. The dynamic nature of cliff quality in the long term is partly explained by the autocorrelation of the prevalence of an ectoparasite that influences reproductive success. As predicted by the performance-based conspecific attraction hypothesis, the reproductive success of current breeders on a given cliff was predictive of the reproductive success of new recruits on the cliff in the following year. Breeders tended to recruit to the previous year’s most productive cliffs and to emigrate from the least productive ones. Consequently, the dynamics of breeder numbers on the cliffs were explained by local reproductive success on a year-to-year basis. Because, on average, young Black-legged Kittiwakes first breed when 4 yr old, such a relationship probably results from individual choices based on the assessment of previous-year local quality. When breeders changed breeding cliffs between years, they selected cliffs of per capita higher reproductive success. Furthermore, after accounting for the potential effects of age and sex as well as between-year variations, the effect of individual breeding performance on breeding dispersal was strongly influenced by the average reproductive success of other breeders on the same cliff. Individual breeding performance did not appear to influence the probability of dispersing for birds breeding on cliffs with high local reproductive success, whereas individual breeding performance did have a strong effect on dispersal for birds that bred on cliffs with lower local reproductive success. This suggests that the reproductive success of locally breeding conspecifics may be sufficient to override an individual’s own breeding experience when deciding whether to emigrate. These results, which are supported by behavioral observations of the role of prospecting in recruitment, suggest that both first breeders and adults rely on the reproductive success of conspecifics as “public information” to assess their own chances of breeding successfully in a given patch and to make settling decisions. A corollary prediction is that individuals should attempt to breed near successful conspecifics (a form of social attraction) in order to benefit from the same favorable local environmental conditions. Such a performance-based conspecific attraction mechanism can thus lead to an aggregative distribution of nests and may have played a role in the evolution of coloniality.

Ecology↗

Expanding national-scale wildlife disease surveillance systems with research networks

Efficient learning about disease dynamics in free-ranging wildlife systems can benefit from active surveillance that is standardized across different ecological contexts. For example, active surveillance that targets specific individuals and populations with standardized sampling across ecological contexts (landscape-scale targeted surveillance) is important for developing a mechanistic understanding of disease emergence, which is the foundation for improving risk assessment of zoonotic or wildlife-livestock disease outbreaks and predicting hotspots of disease emergence. However, landscape-scale targeted surveillance systems are rare and challenging to implement. Increasing experience and infrastructure for landscape-scale targeted surveillance will improve readiness for rapid deployment of this type of surveillance in response to new disease emergence events. Here, we describe our experience developing and rapidly deploying a landscape-scale targeted surveillance system for severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) in two free-ranging deer species across their ranges in the United States. Our surveillance system was designed to collect data across individual, population, and landscape scales for future analyses aimed at understanding mechanisms and risk factors of SARS-CoV-2 transmission, evolution, and persistence. Our approach leveraged partnerships between state and federal public service sectors and academic researchers in a landscape-scale targeted surveillance research network. Methods describe our approach to developing the surveillance network and sampling design. Results report challenges with implementing our intended sampling design, specifically how the design was adapted as different challenges arose and summarize the sampling design that has been implemented thus far. In the discussion, we describe strategies that were important for the successful deployment of landscape-scale targeted surveillance, development and operation of the research network, construction of similar networks in the future, and analytical approaches for the data based on the sampling design.

Ecology and Evolution↗

Interactions between fire and invasive plants in temperate grasslands of North America

A substantial number of invasive grasses, forbs and woody plants have invaded temperate grasslands in North America. Among the invading species are winter annuals, biennials, cool-season perennials, warm-season perennials, vines, shrubs, and trees. Many of these species have been deliberately introduced and widely planted; some are still used for range improvement, pastures, lawns, and as ornamentals, though many are listed as state or federal noxious weeds. Others have been greatly facilitated by widespread land disturbance. Historically, fire has been a major selective force in the evolution of temperate grasslands. Further, prescribed fire is commonly used as a method of ecological management for native grassland communities as well as in conjunction with restoration efforts. Within this context, it is important to understand how invasive species will interact with natural and prescribed fire regimes. In this paper, we consider what is known about how exotic species that invade temperate grasslands relate to fire. The primary issues addressed for each species are (1) Does fire appear to enhance colonization? (2) To what degree does fire affect the survival of plants? (3) Are plants that are burned able to regrow following fire and, if so, how rapidly can they recover? (4) How important is competition with native species to the response to fire? and (5) What effect does an invasive species have on the characteristics of the fire regime? For many species, results are preliminary, incomplete, or inconsistent among studies. For this reason, many of the conclusions drawn for individual species must be considered preliminary. Based on analyses of individual species, a conceptual framework is presented for considering how invasive plants may interact with fire when they invade an ecosystem. The major categories of influences are the native community, the fire regime, growth conditions for both invasive and native species, and influences that disturbances, human impacts, and landscape characteristics have had in the past and will have in the future. The examples considered in this paper provide support for a few, tentative generalizations. First, among our current worst invaders of temperate grasslands, adaptation to fire is quite variable. Some species are not well adapted to burning and can be easily eliminated; other species are better adapted but can still be eliminated if fire occurs during periods of particular vulnerability and/or at high frequency. There is a set of species that is extremely well adapted to fire and will not be eliminated through burning alone. Second, competitive interactions with native species play a crucial role in the success of nonnative invaders. In cases where differential burn responses between invasive and native species can be exploited, and adequate populations of native dominant species are present, fire can sometimes tip the competitive balance away from invasives. Third, there are a few invasive species that have exceptional attributes and for which there are no easy solutions. The ability of cheatgrass ( Bromus tectorum ) to enhance fire, the ability of Chinese tallow ( Triadica sebifera ) to suppress fire, and the ability of leafy spurge ( Euphorbia esula ) to resprout from repeated injury make these species exceptional threats to native diversity. Finally, the available information for many invasive species is very incomplete, partic- ularly with regard to how fire affects competitive interactions with the native community. There is much more we need to know if we are to consistently predict how invasive species will respond to fire and how burning can best be used to manage for natural diversity.

Conference Paper↗

Heightened exposure to parasites favors the evolution of immunity in brood parasitic cowbirds

Immunologists and evolutionary biologists are interested in how the immune system evolves to fit an ecological niche. We studied the relationship between exposure to parasites and strength of immunity by investigating the response of two species of New World cowbirds (genus Molothrus , Icteridae), obligate brood parasites with contrasting life history strategies, to experimental arboviral infection. The South American shiny cowbird ( M. bonariensis ) is an extreme host - generalist that lays its eggs in the nests of >225 different avian species. The Central American bronzed cowbird ( M. aeneus ) is a relative host - specialist that lays its eggs preferentially in the nests of approximately 12 orioles in a single sister genus. West Nile virus provided a strong challenge and delineated immune differences between these species. The extreme host-generalist shiny cowbird, like the North American host-generalist, the brown-headed cowbird, showed significantly lower viremia to three arboviruses than related icterid species that were not brood parasites. The bronzed cowbird showed intermediate viremia. These findings support the interpretation that repeated exposure to a high diversity of parasites favors the evolution of enhanced immunity in brood parasitic cowbirds and makes them useful models for future studies of innate immunity.

Evolutionary Biology↗

Evolution of a reassortant North American gull influenza virus lineage: drift, shift and stability

Background: The role of gulls in the ecology of avian influenza (AI) is different than that of waterfowl. Different constellations of subtypes circulate within the two groups of birds and AI viruses isolated from North American gulls frequently possess reassortant genomes with genetic elements from both North America and Eurasian lineages. A 2008 isolate from a Newfoundland Great Black-backed Gull contained a mix of North American waterfowl, North American gull and Eurasian lineage genes. Methods: We isolated, sequenced and phylogenetically compared avian influenza viruses from 2009 Canadian wild birds. Results: We analyzed six 2009 virus isolates from Canada and found the same phylogenetic lineage had persisted over a larger geographic area, with an expanded host range that included dabbling and diving ducks as well as gulls. All of the 2009 virus isolates contained an internal protein coding set of genes of the same Eurasian lineage genes except PB1 that was from a North American lineage, and these genes continued to evolve by genetic drift. We show evidence that the 2008 Great Black-backed Gull virus was derived from this lineage with a reassortment of a North American PA gene into the more stable core set of internal protein coding genes that has circulated in avian populations for at least 2 years. From this core, the surface glycoprotein genes have switched several times creating H13N6, H13N2, and H16N3 subtypes. These gene segments were from North American lineages except for the H16 and N3 vRNAs. Conclusions: This process appears similar to genetic shifts seen with swine influenza where a stable "triple reassortant internal gene" core has circulated in swine populations with genetic shifts occurring with hemaggluttinin and neuraminidase proteins getting periodically switched. Thus gulls may serve as genetic mixing vessels for different lineages of avian influenza, similar to the role of swine with regards to human influenza. These findings illustrate the need for continued surveillance in gull and waterfowl populations, both on the Pacific and especially Atlantic coasts of North America, to document virus intercontinental movement and the role of gull species in the evolution and epidemiology of AI.

Virology Journal↗

Lineage diversification and morphological evolution in a large-scale continental radiation: The neotropical ovenbirds and woodcreepers (Aves: Furnariidae)

Patterns of diversification in species-rich clades provide insight into the processes that generate biological diversity. We tested different models of lineage and phenotypic diversification in an exceptional continental radiation, the ovenbird family Furnariidae, using the most complete species-level phylogenetic hypothesis produced to date for a major avian clade (97% of 293 species). We found that the Furnariidae exhibit nearly constant rates of lineage accumulation but show evidence of constrained morphological evolution. This pattern of sustained high rates of speciation despite limitations on phenotypic evolution contrasts with the results of most previous studies of evolutionary radiations, which have found a pattern of decelerating diversity-dependent lineage accumulation coupled with decelerating or constrained phenotypic evolution. Our results suggest that lineage accumulation in tropical continental radiations may not be as limited by ecological opportunities as in temperate or island radiations. More studies examining patterns of both lineage and phenotypic diversification are needed to understand the often complex tempo and mode of evolutionary radiations on continents.

Evolution↗

A long-term study of ecological impacts of river channelization on the population of an endangered fish: Lessons learned for assessment and restoration

Projects to assess environmental impact or restoration success in rivers focus on project-specific questions but can also provide valuable insights for future projects. Both restoration actions and impact assessments can become “adaptive” by using the knowledge gained from long-term monitoring and analysis to revise the actions, monitoring, conceptual model, or interpretation of findings so that subsequent actions or assessments are better informed. Assessments of impact or restoration success are especially challenging when the indicators of interest are imperiled species and/or the impacts being addressed are complex. From 1997 to 2015, we worked closely with two federal agencies to monitor habitat availability for and population density of Roanoke logperch ( Percina rex ), an endangered fish, in a 24-km-long segment of the upper Roanoke River, VA. We primarily used a Before-After-Control-Impact analytical framework to assess potential impacts of a river channelization project on the P. rex population. In this paper, we summarize how our extensive monitoring facilitated the evolution of our (a) conceptual understanding of the ecosystem and fish population dynamics; (b) choices of ecological indicators and analytical tools; and (c) conclusions regarding the magnitude, mechanisms, and significance of observed impacts. Our experience with this case study taught us important lessons about how to adaptively develop and conduct a monitoring program, which we believe are broadly applicable to assessments of environmental impact and restoration success in other rivers. In particular, we learned that (a) pre-treatment planning can enhance monitoring effectiveness, help avoid unforeseen pitfalls, and lead to more robust conclusions; (b) developing adaptable conceptual and analytical models early was crucial to organizing our knowledge, guiding our study design, and analyzing our data; (c) catchment-wide processes that we did not monitor, or initially consider, had profound implications for interpreting our findings; and (d) using multiple analytical frameworks, with varying assumptions, led to clearer interpretation of findings than the use of a single framework alone. Broader integration of these guiding principles into monitoring studies, though potentially challenging, could lead to more scientifically defensible assessments of project effects.

Water↗

Nest predation risk explains variation in avian clutch size

Questions about the ecological drivers of, and mechanistic constraints on, productivity have driven research on life-history evolution for decades. Resource availability and offspring mortality are considered among the 2 most important influences on the number of offspring per reproductive attempt. We used a factorial experimental design to manipulate food abundance and perceived offspring predation risk in a wild avian population (red-faced warblers; Cardellina rubrifrons ) to identify the mechanistic cause of variation in avian clutch size. Additionally, we tested whether female quality helped explain the extant variation in clutch size. We found no support for the Food Limitation or Female Quality Hypotheses, but we did find support for both predictions of the Nest Predation Risk Hypothesis. Females that experienced an experimentally heightened perception of offspring predation risk responded by laying a smaller clutch than females in the control group. Additionally, predation rates at artificial nests were highest where red-faced warbler clutch size was smallest (at high elevations). Life-history theory predicts that an individual should invest less in reproduction when high nest predation risk reduces the likely benefit from that nesting attempt and, indeed, we found that birds exhibit phenotypic plasticity in clutch size by laying fewer eggs in response to increasing nest predation risk.

Behavioral Ecology↗

Orthoptera-specific target enrichment (OR-TE) probes resolve relationships over broad phylogenetic scales

Phylogenomic data are revolutionizing the field of insect phylogenetics. One of the most tenable and cost-effective methods of generating phylogenomic data is target enrichment, which has resulted in novel phylogenetic hypotheses and revealed new insights into insect evolution. Orthoptera is the most diverse insect order within polyneoptera and includes many evolutionarily and ecologically interesting species. Still, the order as a whole has lagged behind other major insect orders in terms of transitioning to phylogenomics. In this study, we developed an Orthoptera-specific target enrichment (OR-TE) probe set from 80 transcriptomes across Orthoptera. The probe set targets 1828 loci from genes exhibiting a wide range of evolutionary rates. The utility of this new probe set was validated by generating phylogenomic data from 36 orthopteran species that had not previously been subjected to phylogenomic studies. The OR-TE probe set captured an average of 1037 loci across the tested taxa, resolving relationships across broad phylogenetic scales. Our detailed documentation of the probe design and bioinformatics process is intended to facilitate the widespread adoption of this tool.

Scientific Reports↗

Advances in the hydrogeochemistry and microbiology of acid mine waters

The last decade has witnessed a plethora of research related to the hydrogeochemistry and microbiology of acid mine waters and associated tailings and waste-rock waters. Numerous books, reviews, technical papers, and proceedings have been published that examine the complex bio-geochemical process of sulfide mineral oxidation, develop and apply geochemical models to site characterization, and characterize the microbial ecology of these environments. This review summarizes many of these recent works, and provides references for those investigating this field. Comparisons of measured versus calculated Eh and measured versus calculated pH for water samples from several field sites demonstrate the reliability of some current geochemical models for aqueous speciation and mass balances. Geochemical models are not, however, used to predict accurately time-dependent processes but to improve our understanding of these systems and to constrain possible processes that contribute to actual or potential water quality issues. Microbiological studies are demonstrating that there is much we have yet to learn about the types of different microorganisms and their function and ecology in mine-waste environments. A broad diversity of green algae, bacteria, archaea, yeasts, and fungi are encountered in acid mine waters, and a better understanding of their ecology and function may potentially enhance remediation possibilities as well as our understanding of the evolution of life.

International Geology Review↗

Food web topology and parasites in the pelagic zone of a subarctic lake

Parasites permeate trophic webs with their often complex life cycles, but few studies have included parasitism in food web analyses. Here we provide a highly resolved food web from the pelagic zone of a subarctic lake and explore how the incorporation of parasites alters the topology of the web. 2. Parasites used hosts at all trophic levels and increased both food-chain lengths and the total number of trophic levels. Their inclusion in the network analyses more than doubled the number of links and resulted in an increase in important food-web characteristics such as linkage density and connectance. 3. More than half of the parasite taxa were trophically transmitted, exploiting hosts at multiple trophic levels and thus increasing the degree of omnivory in the trophic web. 4. For trophically transmitted parasites, the number of parasite-host links exhibited a positive correlation with the linkage density of the host species, whereas no such relationship was seen for nontrophically transmitted parasites. Our findings suggest that the linkage density of free-living species affects their exposure to trophically transmitted parasites, which may be more likely to adopt highly connected species as hosts during the evolution of complex life cycles. 5. The study supports a prominent role for parasites in ecological networks and demonstrates that their incorporation may substantially alter considerations of food-web structure and functioning. ?? 2009 British Ecological Society.

Journal of Animal Ecology↗

Stochastic population dynamics in populations of western terrestrial garter snakes with divergent life histories

Comparative evaluations of population dynamics in species with temporal and spatial variation in life‐history traits are rare because they require long‐term demographic time series from multiple populations. We present such an analysis using demographic data collected during the interval 1978–1996 for six populations of western terrestrial garter snakes ( Thamnophis elegans ) from two evolutionarily divergent ecotypes. Three replicate populations from a slow‐living ecotype, found in mountain meadows of northeastern California, were characterized by individuals that develop slowly, mature late, reproduce infrequently with small reproductive effort, and live longer than individuals of three populations of a fast‐living ecotype found at lakeshore locales. We constructed matrix population models for each of the populations based on 8–13 years of data per population and analyzed both deterministic dynamics based on mean annual vital rates and stochastic dynamics incorporating annual variation in vital rates. (1) Contributions of highly variable vital rates to fitness (λ s ) were buffered against the negative effects of stochastic variation, and this relationship was consistent with differences between the meadow (M‐slow) and lakeshore (L‐fast) ecotypes. (2) Annual variation in the proportion of gravid females had the greatest negative effect among all vital rates on λ s . The magnitude of variation in the proportion of gravid females and its effect on λ s was greater in M‐slow than L‐fast populations. (3) Variation in the proportion of gravid females, in turn, depended on annual variation in prey availability, and its effect on λ s was 4–23 times greater in M‐slow than L‐fast populations. In addition to differences in stochastic dynamics between ecotypes, we also found higher mean mortality rates across all age classes in the L‐fast populations. Our results suggest that both deterministic and stochastic selective forces have affected the evolution of divergent life‐history traits in the two ecotypes, which, in turn, affect population dynamics. M‐slow populations have evolved life‐history traits that buffer fitness against direct effects of variation in reproduction and that spread lifetime reproduction across a greater number of reproductive bouts. These results highlight the importance of long‐term demographic and environmental monitoring and of incorporating temporal dynamics into empirical studies of life‐history evolution.

California↗