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U.S. Geological Survey sage-grouse and sagebrush ecosystem research annual report for 2018

The sagebrush ( Artemisia spp.) ecosystem extends across a large portion of the Western United States, and the greater sage-grouse ( Centrocercus urophasianus ) is one of the iconic species of this ecosystem. Greater sage-grouse populations occur in 11 States and are dependent on relatively large expanses of sagebrush-dominated habitat. Sage-grouse populations have been experiencing long-term declines owing to multiple stressors, including interactions among fire, exotic plant invasions, and human land uses, which have resulted in significant loss, fragmentation, and degradation of landscapes once dominated by sagebrush. In addition to the sage-grouse, over 350 species of plants and animals are dependent on the sagebrush ecosystem. Increasing knowledge about how these species and the sagebrush ecosystem respond to these stressors and to management actions can inform and improve strategies to maintain existing areas of intact sagebrush and restore degraded landscapes. The U.S. Geological Survey (USGS) has a broad research program focused on providing the science needed to inform these strategies and to help land and resource managers at the Federal, State, Tribal, and local levels as they work towards sustainable sage-grouse populations and restored landscapes for the broad range of uses critical to stakeholders in the Western United States. USGS science has provided a foundation for major land and resource management decisions including those that precluded the need to list the greater sage-grouse under the Endangered Species Act. The USGS is continuing to build on that foundation to inform science-based decisions to help support local economies and the continued conservation, management, and restoration of the sagebrush ecosystem. This report contains descriptions of USGS sage-grouse and sagebrush ecosystem research projects that are ongoing or were active during 2018 and is organized into five thematic areas: Fire, Invasive Species, Restoration, Sagebrush, Sage-Grouse, and Other Sagebrush-Associated Species; and Climate and Weather.

Circular↗

Fijian sea krait behavior relates to fine-scale environmental heterogeneity in old-growth coastal forest: The importance of integrated land–sea management for protecting amphibious animals

The importance of terrestrial coastal ecosystems for maintaining healthy coral reef ecosystems remains understudied. Sea kraits are amphibious snakes that require healthy coral reefs for foraging, but little is known about their requirements of terrestrial habitats, where they slough their skin, digest prey, and breed. Using concurrent microclimate measurements and behavior surveys, we show that a small, topographically flat atoll in Fiji with coastal forest provides many microhabitats that relate to the behaviors of Yellow Lipped Sea Kraits, Laticauda colubrina . Microclimates were significantly related to canopy cover, leaf litter depth, and distance from the high-water mark (HWM). Sea kraits were almost exclusively observed in coastal forest within 30 m of the HWM. Sloughing of skins only occurred within crevices of mature or dying trees. Resting L. colubrina were significantly more likely to occur at locations with higher mean diurnal temperatures, lower leaf litter depths, and shorter distances from the HWM. On Leleuvia, behavior of L. colubrina therefore relates to environmental heterogeneity created by old-growth coastal forests, particularly canopy cover and crevices in mature and dead tree trunks. The importance of healthy coastal habitats, both terrestrial and marine, for L. colubrina suggests it could be a good flagship species for advocating integrated land-sea management. Furthermore, our study highlights the importance of coastal forests and topographically flat atolls for biodiversity conservation. Effective conservation management of amphibious species that utilize land- and seascapes is therefore likely to require a holistic approach that incorporates connectivity among ecosystems and environmental heterogeneity at all relevant scales.

Ecology and Evolution↗

Multistate models for estimation of survival and reproduction in the Grey-headed Albatross ( Thalassarche chrysostoma )

Reliable information on demography is necessary for conservation of albatrosses, the most threatened family of pelagic birds. Albatross survival has been estimated using mark-recapture data and the Cormack-Jolly-Seber (CJS) model. However, albatross exhibit skipped breeding, violating assumptions of the CJS model. Multistate modeling integrating unobservable states is a promising tool for such situations. We applied multistate models to data on Grey-headed Albatross ( Talassarche chrysostoma ) to evaluate model performance and describe demographic patterns. These included a multistate equivalent of the CJS model (MS-2), including successful and failed breeding states and ignoring temporary emigration, and three versions of a four-state multistate model that accounts for temporary emigration by integrating unobservable states: a model (MS-4) with one sample per breeding season, a robust design model (RDMS-4) with multiple samples per season and geographic closure within the season, and an open robust design model (ORDMS-4) with multiple samples per season and staggered entry and exit of animals within the season. Survival estimates from the MS-2 model were higher than those from the MS-4 model, which resulted in apparent percent relative bias averaging 2.2%. The ORDMS-4 model was more appropriate than the RDMS-4 model, given that staggered entry and exit occurred. Annual survival probability for Greyheaded Albatross at Marion Island was 0.951 ± 0.006 (SE), and the probability of skipped breeding in a subsequent year averaged 0.938 for successful and 0.163 for failed breeders. We recommend that multistate models with unobservable states, combined with robust-design sampling, be used in studies of species that exhibit temporary emigration.

The Auk↗

U.S. Geological Survey sage-grouse and sagebrush ecosystem research annual report for 2017

The sagebrush ( Artemisia spp.) ecosystem extends across a large portion of the Western United States, and the greater sage-grouse ( Centrocercus urophasianus ) is one of the iconic species of this ecosystem. Greater sage-grouse populations occur in 11 States and are dependent on relatively large expanses of sagebrush-dominated habitat. Sage-grouse populations have been experiencing long-term declines owing to multiple stressors, including interactions among fire, exotic plant invasions, and human land uses, which have resulted in significant loss, fragmentation, and degradation of landscapes once dominated by sagebrush. In addition to the sage-grouse, over 350 species of plants and animals are dependent on the sagebrush ecosystem. Increasing knowledge about how these species and the sagebrush ecosystem respond to these stressors and to management actions can inform and improve strategies to maintain existing areas of intact sagebrush and restore degraded landscapes. The U.S. Geological Survey (USGS) has a broad research program focused on providing the science needed to inform these strate-gies and to help land and resource managers at the Federal, State, Tribal, and local levels as they work towards sustainable sage-grouse populations and restored landscapes for the broad range of uses critical to stakeholders in the Western United States. USGS science has provided a foundation for major land and resource management decisions including those that precluded the need to list the greater sage-grouse under the Endangered Species Act. The USGS is continuing to build on that foundation to inform science-based decisions to help support local economies and the continued conservation, management, and restoration of the sagebrush ecosystem. This report contains descriptions of USGS sage-grouse and sagebrush ecosystem research projects that are ongoing or were active during 2017 and is organized into five thematic areas: Fire, Invasive Species, Restoration, Sagebrush and Sage-Grouse, and Climate and Weather.

Circular↗

Hidden Markov movement models reveal diverse seasonal movement patterns in two North American ungulates

Animal movement is the mechanism connecting landscapes to fitness, and understanding variation in seasonal animal movements has benefited from the analysis and categorization of animal displacement. However, seasonal movement patterns can defy classification when movements are highly variable. Hidden Markov movement models (HMMs) are a class of latent-state models well-suited to modeling movement data. Here, we used HMMs to assess seasonal patterns of variation in the movement of pronghorn ( Antilocapra americana ), a species known for variable seasonal movements that challenge analytical approaches, while using a population of mule deer ( Odocoileus hemionus ), for whom seasonal movements are well-documented, as a comparison. We used population-level HMMs in a Bayesian framework to estimate a seasonal trend in the daily probability of transitioning between a short-distance local movement state and a long-distance movement state. The estimated seasonal patterns of movements in mule deer closely aligned with prior work based on indices of animal displacement: a short period of long-distance movements in the fall season and again in the spring, consistent with migrations to and from seasonal ranges. We found seasonal movement patterns for pronghorn were more variable, as a period of long-distance movements in the fall was followed by a winter period in which pronghorn were much more likely to further initiate and remain in a long-distance movement pattern compared with the movement patterns of mule deer. Overall, pronghorn were simply more likely to be in a long-distance movement pattern throughout the year. Hidden Markov movement models provide inference on seasonal movements similar to other methods, while providing a robust framework to understand movement patterns on shorter timescales and for more challenging movement patterns. Hidden Markov movement models can allow a rigorous assessment of the drivers of changes in movement patterns such as extreme weather events and land development, important for management and conservation.

Wyoming↗

Identification of novel hepaciviruses and Sylvilagus-associated viruses via metatranscriptomics in North American lagomorphs

Cottontails ( Sylvilagus spp.) and jackrabbits ( Lepus spp.) within the Leporidae family are native to North America and are found in a wide range of habitats, including deserts, forests, and grasslands. Although there is a growing body of research describing the arrival of the highly virulent rabbit haemorrhagic disease virus 2 (RHDV2, GI.2) on this continent, and its impact on native lagomorphs, information about the natural virome and microbiome of healthy and deceased American lagomorphs is relatively limited. In this study, we used a meta-transcriptomics approach to conduct whole pathogen profiling on healthy and deceased animals in the USA. We analysed 48 matched liver and lung sample pools from apparently healthy cottontails and jackrabbits in Texas and an additional 48 liver samples from deceased animals from nine other US states. This approach enabled the discovery of three distinct new viruses and revealed additional new insights into the lung and liver microbiomes of North American lagomorphs. Of the three new viruses, a tetnovirus and a novel picorna-like virus were likely of insect origin and therefore considered environmental contaminants. Of particular interest was a new species of hepacivirus, with around 50% sequence identity to a known hepacivirus from a xeric four-striped grass rat ( Rhabdomys pumilio ). Phylogenetic analysis from 41 individual hepacivirus genomes recovered from our lagomorph samples revealed two distinct clades, corresponding with different cottontail species. No hepaciviruses were detected in any of the jackrabbit samples. This is the first description of a hepacivirus in lagomorphs. Our findings extend the Hepacivirus genus, provide new insights into its evolution, and describe the first baseline on microbial diversity in North American lagomorphs, an important step towards understanding the role of potential pathogens for population management and conservation.

Arizona, California, Iowa, Massachusetts, Montana,↗

Pit tag application in native freshwater mussels: Case studies across small, medium, and large rivers

Since their first use in the mid-1980s, external passive integrated transponder (PIT) tags have facilitated innovative investigations into multiple biological traits of animals. For native freshwater mussels, PIT tags are frequently used in capture-mark-recapture applications because they allow repeated, noninvasive sampling, are easy to apply, have high retention rates, and have negligible short-term effects on growth and survival. Because of these traits, resource managers and scientists are using PIT-tagged animals to estimate survival and movement of mussels associated with restoration efforts. However, consistency is limited in how PIT tags are affixed, monitored, and reported. Thus, our objectives were to (1) share our collective experiences in PIT tagging mussels across three case studies in small, medium, and large rivers and (2) propose guidelines for tagging and reporting data from PIT tag studies with native freshwater mussels to facilitate comparisons across future studies. The number of studies that have marked mussels with PIT tags has increased over the past 10 years. The ability to detect mussels using PIT tags has substantially advanced research in three areas of mussel ecology: (1) estimating vital rates (e.g., growth and survival), (2) tracking movements and behaviors of captively propagated, wild, and translocated individuals, and (3) improving our understanding of life history traits, such as reproductive timing. Each case study offers insights on tagging methods, tag loss, tag retention, and monitoring frequency across multiple species that range in conservation status from common to rare. We conclude with best-practice guidelines for placing PIT tags on freshwater mussels and a list of variables that could be reported in future studies to facilitate cross-system comparisons.

Freshwater Mollusk Biology and Conservation↗

Human impact on the planet: an earth system science perspective and ethical considerations

The modern Earth Narrative, the scientific story of the 4.5 billion-year natural and human history of the Earth, has emerged from the solid foundation of two factual concepts: Deep (or Geologic) Time and Biological Evolution. spread acceptance of the Earth Narrative is critically important as we begin the third millennium, because it provides a clear understanding of the growing impact of human population growth and associated activities on the Earth System, especially the negative impact on Earth?s biosphere. It is important for humans to realize that we are but one of 4,500 species of mammals that exist on Earth and that we are but one species in the estimated 30 to 100 million species that form the complex biosphere. We also need to recognize that all species exist within the physical limits imposed by the geosphere. We are totally dependent on the biosphere for food, oxygen, and other necessities of life. mans are one of the latest results of biological evolution operating over a long period of Geologic Time. We find ourselves on Earth, after 4.5 billion years of Earth history by chance, not by design. Humans have become so successful at modifying their environment that many of the natural limitations on the expansion of populations of our fellow animals have been overcome by technological and cultural innovations. According to Peter Raven, ?Humans, at a current population of 6 billion [expected to nearly double by 2050], are consuming or wasting about 50 percent of the total net biological productivity on land and 50 percent of the available supply of freshwater. The overwhelming and expanding human presence leaves less and less room in the environment for other biota.? st century will be a pivotal time in the fate of Earth?s biosphere. Whereas human modification of the geosphere will slowly recover over time, human changes to the biosphere are a far more consequential matter? extinction of a species is forever! Will humans effectively use our new knowledge of natural and human history to stop further degradation of Earth?s ecosystems and extinction of its biota? The fate of the biosphere, including humanity, depends on a reaffirmation by all humans of all cultures and religions of the global importance of a planet-wide conservation of the Earth?s biotic heritage. For the world?s religions it means elevation of stewardship of the Earth to a moral imperative and a goal of complete preservation of the Earth?s biotic inheritance, one which is based on a Do No Harm ethic.

Open-File Report↗

A novel member of the family Hepeviridae from cutthroat trout ( Oncorhynchus clarkii )

Beginning in 1988, the Chinook salmon embryo (CHSE-214) cell line was used to isolate a novel virus from spawning adult trout in the state of California, USA. Termed the cutthroat trout ( Oncorhynchus clarkii ) virus (CTV), the small, round virus was not associated with disease, but was subsequently found to be present in an increasing number of trout populations in the western USA, likely by a combination of improved surveillance activities and the shipment of infected eggs to new locations. Here, we report that the full length genome of the 1988 Heenan Lake isolate of CTV consisted of 7269 nucleotides of positive-sense, single-stranded RNA beginning with a 5' untranslated region (UTR), followed by three open reading frames (ORFs), a 3' UTR and ending in a polyA tail. The genome of CTV was similar in size and organization to that of Hepatitis E virus (HEV) with which it shared the highest nucleotide and amino acid sequence identities. Similar to the genomes of human, rodent or avian hepeviruses, ORF 1 encoded a large, non-structural polyprotein that included conserved methyltransferase, protease, helicase and polymerase domains, while ORF 2 encoded the structural capsid protein and ORF 3 the phosphoprotein. Together, our data indicated that CTV was clearly a member of the family Hepeviridae , although the level of amino acid sequence identity with the ORFs of mammalian or avian hepeviruses (13-27%) may be sufficiently low to warrant the creation of a novel genus. We also performed a phylogenetic analysis using a 262. nt region within ORF 1 for 63 isolates of CTV obtained from seven species of trout reared in various geographic locations in the western USA. While the sequences fell into two genetic clades, the overall nucleotide diversity was low (less than 8.4%) and many isolates differed by only 1-2 nucleotides, suggesting an epidemiological link. Finally, we showed that CTV was able to form persistently infected cultures of the CHSE-214 cell line that may have use in research on the biology or treatment of hepevirus infections of humans or other animals.

Virus Research↗

Distribution of an invasive aquatic pathogen (viral hemorrhagic septicemia virus) in the Great Lakes and its relationship to shipping

Viral hemorrhagic septicemia virus (VHSV) is a rhabdovirus found in fish from oceans of the northern hemisphere and freshwaters of Europe. It has caused extensive losses of cultured and wild fish and has become established in the North American Great Lakes. Large die-offs of wild fish in the Great Lakes due to VHSV have alarmed the public and provoked government attention on the introduction and spread of aquatic animal pathogens in freshwaters. We investigated the relations between VHSV dispersion and shipping and boating activity in the Great Lakes by sampling fish and water at sites that were commercial shipping harbors, recreational boating centers, and open shorelines. Fish and water samples were individually analyzed for VHSV using quantitative reverse transcription-polymerase chain reaction (qRT-PCR) and cell culture assays. Of 1,221 fish of 17 species, 55 were VHSV positive with highly varied qRT-PCR titers (1 to 5,950,000 N gene copies). The detections of VHSV in fish and water samples were closely associated and the virus was detected in 21 of 30 sites sampled. The occurrence of VHSV was not related to type of site or shipping related invasion hotspots. Our results indicate that VHSV is widely dispersed in the Great Lakes and is both an enzootic and epizootic pathogen. We demonstrate that pathogen distribution information could be developed quickly and is clearly needed for aquatic ecosystem conservation, management of affected populations, and informed regulation of the worldwide trade of aquatic organisms.

PLoS ONE↗

Permissible Home Range Estimation (PHRE) in restricted habitats: A new algorithm and an evaluation for sea otters

Parametric and nonparametric kernel methods dominate studies of animal home ranges and space use. Most existing methods are unable to incorporate information about the underlying physical environment, leading to poor performance in excluding areas that are not used. Using radio-telemetry data from sea otters, we developed and evaluated a new algorithm for estimating home ranges (hereafter Permissible Home Range Estimation, or “PHRE”) that reflects habitat suitability. We began by transforming sighting locations into relevant landscape features (for sea otters, coastal position and distance from shore). Then, we generated a bivariate kernel probability density function in landscape space and back-transformed this to geographic space in order to define a permissible home range. Compared to two commonly used home range estimation methods, kernel densities and local convex hulls, PHRE better excluded unused areas and required a smaller sample size. Our PHRE method is applicable to species whose ranges are restricted by complex physical boundaries or environmental gradients and will improve understanding of habitat-use requirements and, ultimately, aid in conservation efforts.

PLoS ONE↗

Animal movement models with mechanistic selection functions

A suite of statistical methods are used to study animal movement. Most of these methods treat animal trajectory data in one of three ways: as discrete pro- cesses, as continuous processes, or as point processes. We brie y review each of these approaches and then focus in on the latter. In the context of point processes, so-called resource selection analyses are among the most common way to statis- tically treat animal trajectory data. However, most resource selection analyses provide inference based on approximations of point process models. The forms of these models have been limited to a few types of specications that provide infer- ence about relative resource use and, less commonly, probability of use. For more general spatio-temporal point process models, the most common type of analysis often proceeds with a data augmentation approach that is used to create a binary data set that can be analyzed with conditional logistic regression. We show that the conditional logistic regression likelihood can be generalized to accommodate a variety of alternative specications related to resource selection. We then provide an example of a case where a spatio-temporal point process model coincides with that implied by a mechanistic model for movement expressed as a partial dier- ential equation derived from rst principles of movement. We demonstrate that inference from this form of point process model is intuitive (and could be useful for management and conservation) by analyzing a set of telemetry data from a mountain lion in Colorado, USA, to understand the eects of spatially explicit environmental conditions on movement behavior of this species.

Spatial Statistics↗

Effects of subsidized predators, resource variability, and human population density on desert tortoise populations in the Mojave Desert, USA

Understanding predator–prey relationships can be pivotal in the conservation of species. For 2 decades, desert tortoise Gopherus agassizii populations have declined, yet quantitative evidence regarding the causes of declines is scarce. In 2005, Ft. Irwin National Training Center, California, USA, implemented a translocation project including 2 yr of baseline monitoring of desert tortoises. Unusually high predation on tortoises was observed after translocation occurred. We conducted a retrospective analysis of predation and found that translocation did not affect the probability of predation: translocated, resident, and control tortoises all had similar levels of predation. However, predation rates were higher near human population concentrations, at lower elevation sites, and for smaller tortoises and females. Furthermore, high mortality rates were not limited to the National Training Center. In 2008, elevated mortality (as high as 43%) occurred throughout the listed range of the desert tortoise. Although no temporal prey base data are available for analysis from any of the study sites, we hypothesize that low population levels of typical coyote Canis latrans prey (i.e. jackrabbits Lepus californicus and other small animals) due to drought conditions influenced high predation rates in previous years. Predation may have been exacerbated in areas with high levels of subsidized predators. Many historical reports of increased predation, and our observation of a range-wide pattern, may indicate that high predation rates are more common than generally considered and may impact recovery of the desert tortoise throughout its range.

Endangered Species Research↗

Post-release survival of fallout Newell’s Shearwater fledglings from a rescue and rehabilitation program on Kauai, Hawaii

Light attraction impacts nocturnally active fledgling seabirds worldwide and is a particularly acute problem on Kaua‘i (the northern-most island in the main Hawaiian Island archipelago) for the Critically Endangered Newell’s shearwater Puffinus newelli . The Save Our Shearwaters (SOS) program was created in 1979 to address this issue and to date has recovered and released to sea more than 30500 fledglings. Although the value of the program for animal welfare is clear, as birds cannot simply be left to die, no evaluation exists to inform post-release survival. We used satellite transmitters to track 38 fledglings released by SOS and compared their survival rates (assessed by tag transmission duration) to those of 12 chicks that fledged naturally from the mountains of Kaua‘i. Wild fledglings transmitted longer than SOS birds, and SOS birds with longer rehabilitation periods transmitted for a shorter duration than birds released immediately or rehabilitated for only 1 d. Although transmitter durations from grounded fledglings were shorter (indicating impacts to survivorship), some SOS birds did survive and dispersed out to sea. All surviving birds (wild and SOS) traveled more than 2000 km to the southwest of Kaua‘i, where they concentrated mostly in the North Pacific Equatorial Countercurrent Province, revealing a large-scale annual post-breeding aggregation zone for fledgling Newell’s shearwaters. While there was reduced survival among birds undergoing rehabilitation, SOS remains an important contribution toward the conservation of Newell’s shearwater because a proportion of released birds do indeed survive. However, light attraction, the root cause of fallout, remains a serious unresolved issue on Kaua’i.

Hawaii↗

Seasonal resource selection of a grassland bird in a dynamic landscape: Importance of a heterogeneous landscape

Habitat loss and fragmentation are a growing threat to wildlife, and a better understanding of these landscape processes is needed to mitigate their effects on species populations. Grassland biomes are among the most imperiled ecosystems in the world, and grassland birds are experiencing significant population declines in North America. Understanding how species respond to differences in resource availabilities across spatiotemporal extents is critical to determining animals' distributions. Here, we investigated the relationship of landscape attributes to spatiotemporal distribution of a grassland bird, the ring-necked pheasant ( Phasianus colchicus ), which has experienced population declines in the Midwest, USA. Pheasant declines have been attributed to two anthropogenic stressors, land use change and climate change. In this study, we evaluated the effect of landscape attributes (composition and configuration) on the home-range size and resource selection of pheasants. We used a 95% fixed kernel estimator to estimate home-range size and identified scales at which landscape features influenced home-range sizes. We quantified landscape features within radii of 250, 500, and 1000 m (i.e., local to broader spatial scales) from the home-range center. We also used resource selection functions to predict the home-range placement (second order) and resource selection within home ranges (third order) of pheasants during winter, pre-nesting, and nesting seasons. We developed multi-scale predictions of pheasant resource selection and identified wetlands, grasslands, Conservation Reserve Program (CRP) grasslands, and small grains as land cover types used by pheasants to fulfill their life requirements. Our results indicated home ranges were more likely to be in a landscape with more CRP, wetlands, and grasslands; more connected grasslands; and a greater number of grassland patches. Pheasants also selected heterogeneous landscape and avoided row crops at both orders of selection. Maintaining habitat heterogeneity, by managing landscapes composed of a high proportion of grasslands and CRP surrounded by small grains and wetlands, could enhance the benefits of local management practices for pheasants. Collectively, insights obtained from our study can advance habitat conservation efforts for similar grassland birds and consequently are of broad utility to biologists and wildlife managers.

South Dakota↗

Assessing tiger population dynamics using photographic capture-recapture sampling

Although wide-ranging, elusive, large carnivore species, such as the tiger, are of scientific and conservation interest, rigorous inferences about their population dynamics are scarce because of methodological problems of sampling populations at the required spatial and temporal scales. We report the application of a rigorous, noninvasive method for assessing tiger population dynamics to test model-based predictions about population viability. We obtained photographic capture histories for 74 individual tigers during a nine-year study involving 5725 trap-nights of effort. These data were modeled under a likelihood-based, ?robust design? capture?recapture analytic framework. We explicitly modeled and estimated ecological parameters such as time-specific abundance, density, survival, recruitment, temporary emigration, and transience, using models that incorporated effects of factors such as individual heterogeneity, trap-response, and time on probabilities of photo-capturing tigers. The model estimated a random temporary emigration parameter of =K' =Y' 0.10 ? 0.069 (values are estimated mean ? SE). When scaled to an annual basis, tiger survival rates were estimated at S = 0.77 ? 0.051, and the estimated probability that a newly caught animal was a transient was = 0.18 ? 0.11. During the period when the sampled area was of constant size, the estimated population size Nt varied from 17 ? 1.7 to 31 ? 2.1 tigers, with a geometric mean rate of annual population change estimated as = 1.03 ? 0.020, representing a 3% annual increase. The estimated recruitment of new animals, Bt, varied from 0 ? 3.0 to 14 ? 2.9 tigers. Population density estimates, D, ranged from 7.33 ? 0.8 tigers/100 km2 to 21.73 ? 1.7 tigers/100 km2 during the study. Thus, despite substantial annual losses and temporal variation in recruitment, the tiger density remained at relatively high levels in Nagarahole. Our results are consistent with the hypothesis that protected wild tiger populations can remain healthy despite heavy mortalities because of their inherently high reproductive potential. The ability to model the entire photographic capture history data set and incorporate reduced-parameter models led to estimates of mean annual population change that were sufficiently precise to be useful. This efficient, noninvasive sampling approach can be used to rigorously investigate the population dynamics of tigers and other elusive, rare, wide-ranging animal species in which individuals can be identified from photographs or other means.

Ecology↗

Use of on-site refugia to protect unionid populations from zebra mussel-induced mortality

Protecting unionid populations as zebra mussels spread into inland waterways has relied mainly on relocating at-risk animals into aquaculture facilities. While such relocations are the only viable management technique for some populations, facility availability is limited, leaving many unionids facing extirpation. Another management strategy is in-situ protection either by enhancing natural refugia or by creating managed refugia. We have reviewed all reports of natural refugia and found that refugia for unionids can be found in many areas. There are many habitats where zebra mussel colonization has been limited, or of a temporary nature. Within zebra mussel infested areas, unionid communities continue to survive in some shallow water sites such as estuaries, deltas, and lake-connected wetlands. Managed refugia can be created in areas where natural refugia do not exist. We present a case study on recent efforts to create refugia in an area with rapidly expanding zebra mussel populations. Preliminary analysis of unionid body condition indicates that removal of encrusted zebra mussels only once a year can improve unionid condition factors and decrease mortality. Natural and managed refugia can provide an additional conservation management option in some areas.

Conference Paper↗

Phylogenetic conservatism in plant phenology

Phenological events – defined points in the life cycle of a plant or animal – have been regarded as highly plastic traits, reflecting flexible responses to various environmental cues. The ability of a species to track, via shifts in phenological events, the abiotic environment through time might dictate its vulnerability to future climate change. Understanding the predictors and drivers of phenological change is therefore critical. Here, we evaluated evidence for phylogenetic conservatism – the tendency for closely related spe-cies to share similar ecological and biological attributes – in phenological traits across flowering plants. We aggregated published and unpublished data on timing of first flower and first leaf, encompassing ~4000 species at 23 sites across the Northern Hemisphere. We reconstructed the phylogeny for the setof included species, first, using the software program Phylomatic, and second, from DNA data. We then quantified phylogenetic conservatism in plant phenology within and across sites. We show that more closely related species tend to flower and leaf at similar times. By contrastingmean flowering times within and across sites, however, we illustrate that it is not the time of yearthat is conserved, but rather the phenological responses to a common set of abiotic cues. Our findings suggest that species cannot be treated as statistically independent when modelling phenological responses. Synthesis. Closely related species tend to resemble each other in the timing of their life-history events, a likely product of evolutionarily conser ved responses to environmental cues. The search for the underlying drivers of phenology must therefore account for species’ shared evolutionary histories.

Journal of Ecology↗