Resource selection by the eastern massasauga rattlesnake on managed land in southwestern Michigan

The eastern massasauga rattlesnake (Sistrurus catenatus catenatus) has experienced population declines throughout its range and is now a candidate for federal protection. However, little is known about massasauga habitat selection in Michigan, particularly in actively managed landscapes. Our objectives were to: 1) quantify whether massasaugas in southwestern Michigan select certain vegetation types disproportionately to their availability and 2) quantify whether the vegetation structure associated with snake locations differed between managed (e.g., burning, woody species removal) and unmanaged stands. We implanted radio transmitters in 51 snakes from 2004 to 2005 and 2008 to 2009. We quantified second-order resource selection using compositional analysis, and modeled the effect of habitat management efforts on vegetation using 4 structural variables. All snakes selected cover types disproportionately to their availability (P = 0.001); a ranking matrix ordered the vegetation types, from most to least used, as: early-mid successional deciduous wetland > early-mid successional deciduous upland > developed > late successional mixed lowland forest > late successional deciduous upland forest. We found that snakes in managed areas were associated with greater amounts of dead herbaceous cover (P = 0.005) and less woody stem density (P < 0.001) and tree dominance (P < 0.001) than were snakes in unmanaged areas; however, live herbaceous cover was comparable. Our results can be used by regional managers to provide early and mid successional habitat with structure similar to that selected by snakes in Michigan. © 2011 The Wildlife Society.     

Massasauga Home Range Patterns in the Midwest

Abstract: The massasauga (Sistrurus catenatus) has drastically declined throughout its range mainly due to habitat loss and human persecution. Populations of the massasauga that occur north and east of the Missouri River, USA, are currently candidates for listing under the United States Endangered Species Act. In areas where land managers wish to improve habitat for existing populations or create habitat for repatriation efforts, an understanding of massasauga spatial ecology is necessary to ensure that adequate sized blocks of habitat are created and properly managed. We studied spatial ecology of massasaugas at 2 sites in Wisconsin and 3 sites in Missouri over an 11-year period. Our results indicated that male massasaugas had larger spatial requirements than all other cohorts. Our study sites supporting viable populations indicated that managers interested in restoring or enhancing massasauga habitat should use 100 ha as a minimum restoration or management target. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):754–759; 2008)     

Evaluating Basking-Habitat Deficiency in the Threatened Eastern Massasauga Rattlesnake

ABSTRACT Woody plant succession is hypothesized to threaten many reptile populations by reducing the amount of solar energy available for thermoregulation. Mitigation via vegetation management is often recommended; however, the need for such management practices rarely has been evaluated. We examined the need for basking-site enhancement for the eastern massasauga rattlesnake (Sistrurus c. catenatus; hereafter EMR) in New York, USA, where only 2 populations remain: one at an open-canopy site and another at a closed-canopy site. Microhabitat temperatures were substantially lower at the closed-canopy site, where EMRs selected the warmest available basking sites. Eastern massasauga rattlesnakes in the open-canopy population selected basking sites that afforded greater cryptic cover. We recommend experimental reduction of shrub cover to improve EMR basking habitat at the closed-canopy site. More generally, we caution that management efforts to reduce shrub cover for basking EMRs should maintain adequate cryptic cover.     

Genetic structure of populations of the threatened eastern massasauga rattlesnake, Sistrurus c. catenatus: evidence from microsatellite DNA markers

Throughout its distribution in North America, the threatened eastern massasauga rattlesnake ( Sistrurus c. catenatus ) persists in a series of habitat-isolated disjunct populations of varying size. Here, we use six microsatellite DNA loci to generate information on the degree of genetic differentiation between, and the levels of inbreeding within populations to understand how evolutionary processes operate in these populations and aid the development of conservation plans for this species. Samples were collected from 199 individuals from five populations in Ontario, New York and Ohio. Our results show that all sampled populations: (i) differ significantly in allele frequencies even though some populations are < 50 km apart, and may contain genetically distinct subpopulations < 2 km apart; (ii) have an average of 23% of alleles that are population specific; and (iii) have significant F _IS values (mean overall F _IS= 0.194) probably due to a combination of Wahlund effects resulting from fine-scale genetic differentiation within populations and the presence of null alleles. Our results imply that massasauga populations may be genetically structured on an extremely fine scale even within continuous populations, possibly due to limited dispersal. Additional information is needed to determine if dispersal and mating behaviour within populations can account for this structure and whether the observed differentiation is due to random processes such as drift or to local adaptation. From a conservation perspective, our results imply that these massasauga populations should be managed as demographically independent units and that each has high conservation value in terms of containing unique genetic variation.     

Biogeographic consequences of shifting climate for the western massasauga (Sistrurus tergeminus)

The western massasauga (Sistrurus tergeminus) is a small pit viper with an extensive geographic range, yet observations of this species are relatively rare. They persist in patchy and isolated populations, threatened by habitat destruction and fragmentation, mortality from vehicle collisions, and deliberate extermination. Changing climates may pose an additional stressor on the survival of isolated populations. Here, we evaluate historic, modern, and future geographic projections of suitable climate for S. tergeminus to outline shifts in their potential geographic distribution and inform current and future management. We used maximum entropy modeling to build multiple models of the potential geographic distribution of S. tergeminus. We evaluated the influence of five key decisions made during the modeling process on the resulting geographic projections of the potential distribution, allowing us to identify areas of model robustness and uncertainty. We evaluated models with the area under the receiver operating curve and true skill statistic. We retained 16 models to project both in the past and future multiple general circulation models. At the last glacial maximum, the potential geographic distribution associated with S. tergeminus occurrences had a stronghold in the southern part of its current range and extended further south into Mexico, but by the mid‐Holocene, its modeled potential distribution was similar to its present‐day potential distribution. Under future model projections, the potential distribution of S. tergeminus moves north, with the strongest northward trends predicted under a climate scenario increase of 8.5 W/m². Some southern populations of S. tergeminus have likely already been extirpated and will continue to be threatened by shifting availability of suitable climate, as they are already under threat from desertification of grasslands. Land use and habitat loss at the northern edge of the species range are likely to make it challenging for this species to track suitable climates northward over time.     

Drift,selection and adaptive variation in small populations of a threatened rattlesnake

An important goal of conservation genetics is to determine if the viability of small populations is reduced by a loss of adaptive variation due to genetic drift. Here, we assessed the impact of drift and selection on direct measures of adaptive variation (toxin loci encoding venom proteins) in the eastern massasauga rattlesnake (Sistrurus catenatus), a threatened reptile that exists in small isolated populations. We estimated levels of individual polymorphism in 46 toxin loci and 1,467 control loci across 12 populations of this species, and compared the results with patterns of selection on the same loci following speciation of S. catenatus and its closest relative, the western massasauga (S. tergeminus). Multiple lines of evidence suggest that both drift and selection have had observable impacts on standing adaptive variation. In support of drift effects, we found little evidence for selection on toxin variation within populations and a significant positive relationship between current levels of adaptive variation and long‐ and short‐term estimates of effective population size. However, we also observed levels of directional selection on toxin loci among populations that are broadly similar to patterns predicted from interspecific selection analyses that pre‐date the effects of recent drift, and that functional variation in these loci persists despite small short‐term effective sizes. This suggests that much of the adaptive variation present in populations may represent an example of “drift debt,” a nonequilibrium state where present‐day levels of variation overestimate the amount of functional genetic diversity present in future populations.     

Similarity of contemporary and historical gene flow among highly fragmented populations of an endangered rattlesnake

Populations of endangered taxa in recently fragmented habitats often show high levels of genetic structure, but the role that contemporary versus historical processes play in generating this pattern is unclear. The eastern massasauga rattlesnake (Sistrurus c. catenatus) is an endangered snake that presently occurs throughout central and eastern North America in a series of populations that are isolated because of habitat fragmentation and destruction. Here, we use data from 19 species-specific microsatellite DNA loci to assess the levels of genetic differentiation, genetic effective population size, and contemporary and historical levels of gene flow for 19 populations sampled across the range of this snake. Eastern massasaugas display high levels of genetic differentiation (overall θ(Fst) = 0.21) and a Bayesian clustering method indicates that each population represents a unique genetic cluster even at regional spatial scales. There is a twofold variation in genetically effective population sizes but little genetic evidence that populations have undergone recent or historical declines in size. Finally, both contemporary and historical migration rates among populations were low and similar in magnitude even for populations located <7 km apart. A test of alternate models of population history strongly favours a model of long-term drift-migration equilibrium over a recent isolation drift-only model. These results suggest that recent habitat fragmentation has had little effect on the genetic characteristics of these snakes, but rather that this species has historically existed in small isolated populations that may be resistant to the long-term negative effects of inbreeding.     

Genetic analysis of populations of threatened snake species using RAPD markers

Snakes are a particularly threatened vertebrate taxon, with distributions of many species and populations becoming increasingly fragmented. At present, little is known about the degree of genetic differentiation that exists between isolated populations even though such information may be critical to their survival and conservation. As an example of how recently developed RAPD genetic markers can be used in conservation genetics, we present preliminary results from a study which used these DNA-based markers to assess population divergence in two threatened Canadian snakes, the black rat snake ( Elaphe o. obsoleta ) and the eastern massasauga rattlesnake ( Sistrurus c. catenatus ). We present information on the levels of variation and reliability of amplification for fragments generated from five primers. We then use a recently developed analytical technique to estimate levels of nucleotide diversity within populations and sequence divergence between populations. Our results show that intrapopulation levels of divergence as estimated by the methods of Clark & Lanigan ( Molecular Biology and Evolution 1993, 10 , 1096–1111) approximate those found for mtDNA in vertebrates and that diversity between snake populations is small and non-significant when tested using randomization procedures. Thus, our study provides an example of how RAPDs can be applied to conservation genetic studies of vertebrates and suggest that the snake populations we examined have only recently become isolated and maybe considered genetically equivalent from a conservation perspective, although this conclusion needs to be confirmed with other DNA-based markers.     

Effects of Habitat on Mallard Duckling Survival in the Great Lakes Region

Abstract: Habitat provides food and shelter resources for prefledgling waterfowl and thus plays a critical role in their growth, development, and survival. However, few studies have examined whether and how particular elements of habitat affect duckling survival. We investigated relationships of duckling survival rates with distance of overland travel, wetland vegetation composition, water permanency, and surrounding upland vegetation for 116 mallard (Anas platyrhynchos) broods in the Great Lakes region from 2001 to 2003. We found that the probability, on hatch day, that a mallard duckling will survive to 55 days was positively related to the proportion of wetland area that was vegetated and negatively related to the proportion of forest cover within 500 m of duckling locations. We found little support for relationships between duckling survival rates and the proportions of grasslands or seasonal wetlands or to distances traveled overland by broods. Our results suggest that conservation groups and wildlife managers in the Great Lakes region can improve mallard duckling survival rates by managing for, creating, and protecting vegetated wetlands and focusing efforts within lightly-forested areas.     

Abundance and distribution of picoplankton in tropical, oligotrophic Lake Kivu, eastern Africa

1. We used flow cytometry to characterize freshwater photosynthetic picoplankton (PPP) and heterotrophic bacteria (HB) in Lake Kivu, one of the East‐African great lakes. Throughout three cruises run in different seasons, covering the four major basins, phycoerythrin‐rich cells dominated the PPP. Heterotrophic bacteria and PPP cell numbers were always high and spatial variations were modest. This represents an important difference from temperate and high latitude lakes that show high fluctuations in cell abundance over an annual cycle. 2. Three populations of picocyanobacteria were identified: one corresponded to single‐cells (identified as Synechococcus by epifluorescence microscopy, molecular methods and pigment content), and the two other that most probably correspond to two and four celled colonies of the same taxon. The proportion of these two subpopulations was greater under stratified conditions, with stronger nutrient limitation. 3. High PPP concentrations (c. 10⁵ cell mL^?1) relative to HB (c. 10⁶ cell mL^?1) were always found. Lake Kivu supports relatively less bacteria than phytoplankton biomass than temperate systems, probably as a consequence of factors such as temperature, oligotrophy, nutrient limitation and trophic structure. 4. A review of PPP concentration across aquatic systems suggests that the abundance of Synechococcus‐like cyanobacteria in large, oligotrophic, tropical lakes is very high. 5. Photosynthetic picoplankton cell abundances in the oligotrophic tropical lakes Kivu and Tanganyika are comparable to those of eutrophic temperate lakes. This apparently contradicts the view that PPP abundance increases with increasing eutrophy. More data on PPP in tropical lakes are needed to explore further this particular pattern.     

Proximate Cues for a Short-Distance Migratory Species: an Application of Survival Analysis

ABSTRACT Investigation of bird migration has often highlighted the importance of external factors in determining timing of migration. However, little distinction has been made between short- and long-distance migrants and between local and flight birds (passage migrants) in describing migration chronology. In addition, measures of food abundance as a proximate factor influencing timing of migration are lacking in studies of migration chronology. To address the relationship between environmental variables and timing of migration, we quantified the relative importance of proximate external factors on migration chronology of local American woodcock (Scolopax minor), a short distance migrant, using event-time analysis methods (survival analysis). We captured 1,094 woodcock local to our study sites in Michigan, Minnesota, and Wisconsin (USA) during autumn 2002–2004 and documented 786 departure dates for these birds. Photoperiod appeared to provide an initial proximate cue for timing of departure. Moon phase was important in modifying timing of departure, which may serve as a navigational aid in piloting and possibly orientation. Local synoptic weather variables also contributed to timing of departure by changing the rate of departure from our study sites. We found no evidence that food availability influenced timing of woodcock departure. Our results suggest that woodcock use a conservative photoperiod-controlled strategy with proximate modifiers for timing of migration rather than relying on abundance of their primary food, earthworms. Managing harvest pressure on local birds by adjusting season lengths may be an effective management tool with consistent migration patterns from year to year based on photoperiod.     

Complex postglacial recolonization inferred from population genetic structure of mottled sculpin Cottus bairdii in tributaries of eastern Lake Michigan,U.S.A.

This study used analyses of the genetic structure of a non‐game fish species, the mottled sculpin Cottus bairdii to hypothesize probable recolonization routes used by cottids and possibly other Laurentian Great Lakes fishes following glacial recession. Based on samples from 16 small streams in five major Lake Michigan, U.S.A., tributary basins, significant interpopulation differentiation was documented (overall F_ST = 0·235). Differentiation was complex, however, with unexpectedly high genetic similarity among basins as well as occasionally strong differentiation within basins, despite relatively close geographic proximity of populations. Genetic dissimilarities were identified between eastern and western populations within river basins, with similarities existing between eastern and western populations across basins. Given such patterns, recolonization is hypothesized to have occurred on three occasions from more than one glacial refugium, with a secondary vicariant event resulting from reduction in the water level of ancestral Lake Michigan. By studying the phylogeography of a small, non‐game fish species, this study provides insight into recolonization dynamics of the region that could be difficult to infer from game species that are often broadly dispersed by humans.     

Factors influencing productivity of eastern wild turkeys in northeastern South Dakota

Population growth is highly sensitive to changes in reproductive rates for many avian species. Understanding how reproductive rates are related to environmental conditions can give managers insight into factors contributing to population change. Harvest trends of eastern wild turkey in northeastern South Dakota suggest a decline in abundance. We investigated factors influencing reproductive success of this important game bird to identify potential factors contributing to the decline. We monitored nesting rate, nest survival, renesting rate, clutch size, hatchability, and poult survival of 116 eastern wild turkey hens using VHF radio transmitters during the springs and summers of 2017 and 2018. Heavier hens were more likely to attempt to nest than lighter hens, and adult hens were more likely to renest than yearling hens. Nest survival probability was lowest in agricultural fields relative to all other cover types and positively related to horizontal visual obstruction and distance to the nearest road. Daily nest survival probability demonstrated an interaction between temperature and precipitation, such that nest survival probability was lower on warm, wet days, but lowest on dry days. Egg predation was the leading cause of nest failure, followed by haying of the nest bowl and death of the incubating hen. Poults reared by adult hens had a greater probability of survival than poults reared by yearling hens. Our estimate of survival probability of poults raised by yearling hens was low relative to other studies, which may be contributing to the apparent regional population decline. However, there is little managers can do to influence poult survival in yearling hens. Alternatively, we found nest survival probability was lowest for nests initiated in agricultural fields. Wildlife‐friendly harvesting practices such as delayed haying or installation of flushing bars could help increase productivity of eastern wild turkey in northeastern South Dakota.     

Seasonal and interannual effects of hypoxia on fish habitat quality in central Lake Erie

1. Hypoxia occurs seasonally in many stratified coastal marine and freshwater ecosystems when bottom dissolved oxygen (DO) concentrations are depleted below 2–3 mg O₂ L^?1. 2. We evaluated the effects of hypoxia on fish habitat quality in the central basin of Lake Erie from 1987 to 2005, using bioenergetic growth rate potential (GRP) as a proxy for habitat quality. We compared the effect of hypoxia on habitat quality of (i) rainbow smelt, Osmerus mordax mordax Mitchill (young‐of‐year, YOY, and adult), a cold‐water planktivore, (ii) emerald shiner, Notropis atherinoides Rafinesque (adult), a warm‐water planktivore, (iii) yellow perch, Perca flavescens Mitchill (YOY and adult), a cool‐water benthopelagic omnivore and (iv) round goby Neogobius melanostomus Pallas (adult) a eurythermal benthivore. Annual thermal and DO profiles were generated from 1D thermal and DO hydrodynamics models developed for Lake Erie’s central basin. 3. Hypoxia occurred annually, typically from mid‐July to mid‐October, which spatially and temporally overlaps with otherwise high benthic habitat quality. Hypoxia reduced the habitat quality across fish species and life stages, but the magnitude of the reduction varied both among and within species because of the differences in tolerance to low DO levels and warm‐water temperatures. 4. Across years, trends in habitat quality mirrored trends in phosphorus concentration and water column oxygen demand in central Lake Erie. The per cent reduction in habitat quality owing to hypoxia was greatest for adult rainbow smelt and round goby (mean: ?35%), followed by adult emerald shiner (mean: ?12%), YOY rainbow smelt (mean: ?10%) and YOY and adult yellow perch (mean: ?8.5%). 5. Our results highlight the importance of differential spatiotemporally interactive effects of DO and temperature on relative fish habitat quality and quantity. These effects have the potential to influence the performance of individual fish species as well as population dynamics, trophic interactions and fish community structure.     

A statistical analysis of subsampling and an evaluation of the Folsom plankton splitter

Subsampling techniques are important for the determination of precise plankton density estimates. A binomial model of random subsampling, and its Poisson extension, were developed for the purpose of evaluating the performance of compartment-type plankton subsamplers. Two approaches were used to assess the performance of the Folsom plankton splitter on an extensive series of nearshore Lake Michigan crustacean zooplankton samples collected between 1974 and 1979. First, Folsom subsamples were observed to be significantly (p < 0.05) more variable than expected from the random model of subsampling. Second, a random effects ANOVA model was used to compare fractions of the total variance in density estimates that were attributable to subsampling and sampling phases of a specially designed study. Departures from randomness in subsampling were sufficiently small that an analysis of optimal allocation of effort between subsampling and sampling phases, based on the ANOVA model, indicated that only one to three subsamples needed to be examined per sample.     

Effectiveness of Short-Distance Translocation and its Effects on Western Rattlesnakes

ABSTRACT Short-distance translocation (SDT) is commonly used to mitigate snake-human interactions, yet little is known about its effectiveness or its effects on behavior and welfare of snakes. Between April 2004 and October 2005, we evaluated SDT as a conservation and management tool by investigating how 500-m SDT affected spatial ecology, body condition, and behavior of western rattlesnakes (Crotalus oreganus) surgically implanted with radiotransmitters in a field study near Osoyoos, British Columbia, Canada. Of 14 rattlesnakes subjected to SDT, 12 (85.7%) returned on ≥1 occasion (range 1–7 times) to the general area they were removed from. Rattlesnakes that underwent SDT showed an increase in total distance moved over an active season compared to non-translocated snakes, but there was no evidence to suggest SDT had an effect on activity range size. There was no evidence to suggest SDT affected body condition, behavior, or mortality rates. Short-distance translocation to high-quality undisturbed habitats was unsuccessful as a long-term solution to snake-human conflict because most translocated snakes returned to conflict areas within a short time (x̄x = 19.9 ± 8.7 days). However, SDT may be an effective short-term tool to manage snake-human conflict in areas where human presence is seasonal or short-lived if careful attention is paid to species-specific biological needs, habitat quality at the release site, and the location of the release site in relation to conflict areas.     

Postfledging survival of eastern bluebirds in an urbanized landscape

Golf courses ostensibly offer green space in urbanized areas, but it is unclear how suitable these human-modified habitats are for wildlife populations. Golf courses are home to a variety of wildlife, but in particular they have been the focus of research on avian responses to urbanization. Although numerous reproductive and diversity studies have been conducted on birds of golf courses, no research exists on postfledging survival in this created landscape. In 2008 and 2009, we estimated survival of eastern bluebird (Sialia sialis) fledglings using radio telemetry on golf course and other developed sites in Williamsburg, Virginia. We used nest survival models in Program MARK set in an information theoretic framework to assess whether the golf course habitat predicted mortality along with other previously studied variables, such as fledgling age, year, site, body condition, fledging date, and transmitter weight. We found no evidence that inhabiting a golf course increased mortality during the fledgling period, but we did find support for both fledgling age and fledging date as predictors of survival. Mortality decreased for older fledglings and those that fledged later in the season. Cause-specific postfledging survival rates did not differ among sites. Fledgling bluebirds did, however, move into habitat that was significantly more forested and less grassy than their natal habitat. For managers of wildlife on golf courses and other urbanized sites, our study is the first to show that placing nest boxes in manicured habitat may attract birds to areas without suitable habitat for fledglings. © 2011 The Wildlife Society.