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The effect of group size on reproduction in cooperatively breeding gray wolves depends on density

In cooperatively breeding species, large group size is often positively related to reproductive success and group persistence. We have a poor understanding, however, of how group sizes within a population affect reproduction particularly as density varies. We hypothesized that at low densities, wolves in both small and large groups would have similar reproductive rates. At high densities, however, wolves in small groups would have lower reproductive rates compared to those in large groups. Using empirical data from radio-collared wolves in Idaho and Yellowstone National Park, WY, USA (1996–2012), we compared reproductive rates (i.e. proportion reproducing, litter size, pup survival) among small and large groups of wolves as density fluctuated within the populations. Reproductive rates were generally lower for individuals in small groups compared to those in large groups, particularly as density increased. Pup survival, however, was slightly higher for wolves in small groups compared to large groups except at very high densities. Polygamy increased with density regardless of group size, suggesting a polygamy threshold for wolves. Large group size resulted in less parturition failure, more breeding females per group, larger litter sizes, and ultimately more pups recruited per group. Large group size appears advantageous for several, but not all, aspects of reproduction particularly when population density is high.

Animal Conservation

Temperature and winter duration requirements for reproductive success in johnny darter Etheostoma nigrum in the South Platte River basin, Colorado

Changes in water temperature and its seasonal timing influences the physiological processes of many aquatic ectotherms. Wastewater treatment plants (WWTP) contribute to warmer and more consistent water temperatures in streams draining the North American Great Plains, particularly during winter months. Reduced variation in seasonal water temperature may adversely affect fishes that rely on temperature fluctuation or specific overwintering temperatures for reproductive cues. We used johnny darter Etheostoma nigrum (Family Percidae), a thermally sensitive species native to the South Platte River Basin, as a focal species to assess the effects of winter water temperature regulations in Colorado. Our objective was to evaluate the effects of winter stream temperature and winter duration on reproductive success of johnny darter in the laboratory to inform development of water quality criteria for the South Platte River Basin. Winter duration (60, 90 or 120 days) and temperature (4 or 12°C) were crossed in a factorial design of six treatments that simulated warmed effluent-impacted streams as well as streams with a more natural thermal regime. We defined reproductive success in johnny darter by examining four reproductive endpoints: (1) spawning timing, (2) egg and larvae production, (3) egg success, and (4) egg development. Across the four endpoints, seven main response variables were measured: spawning initiation date, eggs per gram of female per week, live larvae per gram of female per week, fertilisation success, hatching success, egg-to-larva survival and days to hatch. Winter temperature and duration influenced spawning timing and egg development. Earlier spawning initiation dates were observed in fish exposed to 12°C winters and was associated with a higher number of days to hatch. Eggs per gram of female per week, live larvae per gram of female per week, fertilisation success, hatching success, and egg-to-larva survival were similar among winter treatments. Although the State of Colorado chronic winter water temperature standard of 12°C appears to be adequate for egg and larvae production in johnny darter, our results showed that fish exposed to the standard began spawning significantly earlier than fish exposed to 4°C winters. Evaluating reproductive output in the context of seasonal timing is valid because spawning timing has the potential to affect overall production and egg development, and consequently recruitment and population sustainability. Therefore, spawning timing should be considered when making management decisions and revising water temperature standards. The warming of streams and rivers from WWTP effluent is a global issue, especially in areas that are undergoing rapid urbanisation. Fishes are particularly sensitive to water temperature and many fishes worldwide are likely to be experiencing changes in important physiological processes, affecting both reproduction and survival. Our study indicates that reproductive timing could be influenced by stream warming and stresses the need for further studies on the influence of anthropogenic effects on stream warming.

Colorado

MASTREE+: Time-series of plant reproductive effort from six continents

Significant gaps remain in understanding the response of plant reproduction to environmental change. This is partly because measuring reproduction in long-lived plants requires direct observation over many years and such datasets have rarely been made publicly available. Here we introduce MASTREE+, a data set that collates reproductive time-series data from across the globe and makes these data freely available to the community. MASTREE+ includes 73,828 georeferenced observations of annual reproduction (e.g. seed and fruit counts) in perennial plant populations worldwide. These observations consist of 5971 population-level time-series from 974 species in 66 countries. The mean and median time-series length is 12.4 and 10 years respectively, and the data set includes 1122 series that extend over at least two decades (≥20 years of observations). For a subset of well-studied species, MASTREE+ includes extensive replication of time-series across geographical and climatic gradients. Here we describe the open-access data set, available as a.csv file, and we introduce an associated web-based app for data exploration. MASTREE+ will provide the basis for improved understanding of the response of long-lived plant reproduction to environmental change. Additionally, MASTREE+ will enable investigation of the ecology and evolution of reproductive strategies in perennial plants, and the role of plant reproduction as a driver of ecosystem dynamics.

Global Change Biology

Reproductive failure of the red shiner (Cyprinella lutrensis) after exposure to an exogenous estrogen

Endocrine disrupting chemicals (EDCs) have been detected in surface waters worldwide and can lead to developmental and reproductive disruption in exposed fishes. In the US Great Plains, EDCs are impacting streams and rivers and may be causing adverse reproductive effects. To examine how estrogenic EDCs might affect reproductive success of plains fishes, we experimentally exposed male red shiners (Cyprinella lutrensis) to exogenous 17b-estradiol. We characterized the effects of estradiol on male gonadal histology and secondary sexual characteristics, determined whether exposure reduced reproductive success, and examined the effects of depuration. Adults were exposed to a mean concentration of 70 ng L-1 estradiol, a solvent control, or a water control for at least 83 days. Male exposure to estradiol resulted in elevated plasma vitellogenin concentrations, changes in spermatogenesis, reduced mating coloration and tubercles, altered mating behaviors, and reduced reproductive success with no viable progeny produced. Reproductive endpoints improved upon depuration (28 days). Exposure to estradiol had significant adverse effects on red shiners, indicating that wild populations may face developmental and reproductive difficulties if they are chronically exposed to estradiol.

Canadian Journal of Fisheries and Aquatic Sciences

Determinants of reproductive costs in the long-lived Black-legged Kittiwake: A multiyear experiment

We studied reproductive costs of Black-legged Kittiwakes ( Rissa tridactyla ) in Prince William Sound, Alaska (USA) by removing entire clutches from randomly selected nests over four successive years, and then contrasting survival and fecundity of adults from manipulated and unmanipulated nests in each subsequent year. To elucidate mechanisms that lead to the expression of reproductive costs, we simultaneously characterized several behavioral and physiological parameters among adults in the two treatment groups. We also examined naturally nonbreeding adults that previously bred to determine their survival and future nonbreeding probabilities. Food availability varied during the study, being generally poor at the onset, and improving in later years. Adult nest attendance and body condition (assessed late in the chick- rearing period) varied accordingly among years, and between adults raising chicks and adults that had their eggs removed. Adults from unmanipulated nests incurred significant survival costs in all years, although fecundity costs were strongly expressed in only one of four years. Neither survival nor fecundity costs were strongly influenced by body condition or food availability, and no difference in reproductive costs was detected between the sexes. Although unmanipulated breeders survived at lower rates than manipulated breeders due to costs of reproduction, their survival rates were elevated compared to those of natural nonbreeders, presumably due to differences in individual ability. These findings indicate that models of adult survival must consider not only an organism's reproductive state, but also the factors that lead to that state. Although body condition appeared to be weakly related to survival, it was insufficient to explain the full magnitude of survival costs observed. We suggest that other parameters that were found to differ between treatment groups (e.g., rates of energy turnover, baseline levels of stress, and patterns of allocating body reserves) may be important mechanistic determinants of reproductive costs in kittiwakes, and potentially other long-lived species. Future efforts should move beyond simple assessments of body condition and toward more integrated measures of physiological condition when attempting to identify factors that influence how long-lived species balance the costs and benefits of reproduction.

Alaska

Estimating breeding proportions and testing hypotheses about costs of reproduction with capture-recapture data

The proportion of animals in a population that breeds is an important determinant of population growth rate. Usual estimates of this quantity from field sampling data assume that the probability of appearing in the capture or count statistic is the same for animals that do and do not breed. A similar assumption is required by most existing methods used to test ecologically interesting hypotheses about reproductive costs using field sampling data. However, in many field sampling situations breeding and nonbreeding animals are likely to exhibit different probabilities of being seen or caught. In this paper, we propose the use of multistate capture-recapture models for these estimation and testing problems. This methodology permits a formal test of the hypothesis of equal capture/sighting probabilities for breeding and nonbreeding individuals. Two estimators of breeding proportion (and associated standard errors) are presented, one for the case of equal capture probabilities and one for the case of unequal capture probabilities. The multistate modeling framework also yields formal tests of hypotheses about reproductive costs to future reproduction or survival or both fitness components. The general methodology is illustrated using capture-recapture data on female meadow voles, Microtus pennsylvanicus. Resulting estimates of the proportion of reproductively active females showed strong seasonal variation, as expected, with low breeding proportions in midwinter. We found no evidence of reproductive costs extracted in subsequent survival or reproduction. We believe that this methodological framework has wide application to problems in animal ecology concerning breeding proportions and phenotypic reproductive costs.

Ecology

Chilled frogs are hot: hibernation and reproduction of the Endangered mountain yellow-legged frog Rana muscosa

In the face of the sixth great extinction crisis, it is imperative to establish effective breeding protocols for amphibian conservation breeding programs. Captive efforts should not proceed by trial and error, nor should they jump prematurely to assisted reproduction techniques, which can be invasive, difficult, costly, and, at times, counterproductive. Instead, conservation practitioners should first look to nature for guidance, and replicate key conditions found in nature in the captive environment, according to the ecological and behavioral requirements of the species. We tested the effect of a natural hibernation regime on reproductive behaviors and body condition in the Endangered mountain yellow-legged frog Rana muscosa . Hibernation had a clear positive effect on reproductive behavior, manifesting in vocal advertisement signaling, female receptivity, amplexus, and oviposition. These behaviors are critical components of courtship that lead to successful reproduction. Our main finding was that captive R. muscosa require a hibernation period for successful reproduction, as only hibernated females produced eggs and only hibernated males successfully fertilized eggs. Although hibernation also resulted in a reduced body condition, the reduction appeared to be minimal with no associated mortality. The importance of hibernation for reproduction is not surprising, since it is a major component of the conditions that R. muscosa experiences in the wild. Other amphibian conservation breeding programs can also benefit from a scientific approach that tests the effect of natural ecological conditions on reproduction. This will ensure that captive colonies maximize their role in providing genetic reservoirs for assurance and reintroduction efforts.

Endangered Species Research

Reproductive frequency and size-dependence of fecundity in the Giant Gartersnake (Thamnophis gigas)

How reproductive output changes with age or size is a key life-history trait that can affect which demographic rates most influence population growth. Although many studies have investigated the reproductive ecology of gartersnakes, we know little about reproduction in the threatened Giant Gartersnake, Thamnophis gigas . We used X-radiography to determine reproductive status and estimated fecundity for 73 female T. gigas collected from several regions within the range of this species in the Sacramento Valley of California, USA, and synthesize these data with data from litters born in captivity to improve our understanding of reproduction in this species. Average total litter size determined from X-rays (15.9) and captive-born litters (15.5) are within the ranges reported from other gartersnakes, but captive-born litters had high rates of stillbirth. Only 154 of 202 neonates from captive snakes were born alive, and seven of 13 litters contained at least one stillborn neonate. We found that fecundity was positively related to maternal snout-vent length, and some evidence that larger litters contained smaller neonates. The proportion of X-rayed females that were gravid was 0.50 in 2014, 0.47 in and 2015, and 0.64 in 2016. Central California experienced an exceptional drought from 2012–2015, which may have affected the reproductive output and frequency of T. gigas . Our estimates of reproductive frequency and size-dependent fecundity in T. gigas provide valuable information that can be used in demographic models of this threatened species. Our results demonstrate that X-radiography is a useful, minimally invasive means to study fecundity in wild populations of snakes.

California

Reproduction partially compensates for human-caused mortality in a cooperative breeder

Reproductive output can vary widely among mammalian species. There are many drivers that affect reproductive output including evolutionary, environmental, population, social, and individual traits. Although several factors, including human-caused mortality, can affect reproductive output, we generally have a poor understanding of how such factors interact to affect reproduction, particularly in cooperative breeders. Gray wolves ( Canis lupus ) in Idaho, USA, are exposed to annual hunting and trapping. Thus, they are an ideal species to answer questions about how turnover within groups affects reproduction in cooperative breeders. I hypothesized that the reproductive output of wolves would be affected by individual, social, and environmental factors. Contrary to my prediction, mid-summer litter size was positively associated with wolf harvest density, suggesting a compensatory response to harvest in cooperatively breeding gray wolves. Such compensation is only partial, however, and does not fully account for all the individuals lost from harvest. At the very highest harvest densities observed, mean litter size increased nearly 28%. In contrast, mid-summer litter size was negatively associated with multiple breeding in groups, suggesting resource limitation and competition within groups. I show that characteristics associated with harvest and breeding strategies predict variations in litter size in a cooperative breeder.

Idaho

Alternate reproductive strategies in the California gull

We analysed 6 years of reproduction data for 176 California gulls ( Larus californicus ) surviving from 1980 to 1988. Using a statistical model adapted from Rao's (1958) and Tucker's (1966) generalized growth curve analysis, we reconstructed the reproductive patterns of gulls aged from 0 to 26 years. Individuals were highly consistent in following one of two patterns of reproduction. In a primary pattern employed by most gulls, individuals skipped breeding less frequently and laid larger clutches as they aged. Clutch size increased to a plateau and remained at high levels throughout remaining life. In an alternate pattern employed by a smaller subset of the sample, clutch size also increased to a plateau. However, as a result of frequent skipping of breeding and smaller clutches, this plateau was considerably lower compared to that of gulls adopting the primary reproductive pattern. Data on fledging success from 1980 and 1984 were consistent with the finding of two reproductive patterns. Gulls adopting the alternate reproductive pattern produce fewer offspring per breeding attempt but survive longer than gulls adopting the primary pattern. The frequency of gulls employing the alternate pattern will increase with age relative to gulls employing the primary pattern. The alternate pattern, and not senescence, may explain why several cross-sectional studies on seabirds report declines among the oldest breeders in measures of clutch size, egg mass, hatching success, and fledging success.

Wyoming

Scale-dependent feedbacks between patch size and plant reproduction in desert grassland

Theoretical models suggest that scale-dependent feedbacks between plant reproductive success and plant patch size govern transitions from highly to sparsely vegetated states in drylands, yet there is scant empirical evidence for these mechanisms. Scale-dependent feedback models suggest that an optimal patch size exists for growth and reproduction of plants and that a threshold patch organization exists below which positive feedbacks between vegetation and resources can break down, leading to critical transitions. We examined the relationship between patch size and plant reproduction using an experiment in a Chihuahuan Desert grassland. We tested the hypothesis that reproductive effort and success of a dominant grass ( Bouteloua eriopoda ) would vary predictably with patch size. We found that focal plants in medium-sized patches featured higher rates of grass reproductive success than when plants occupied either large patch interiors or small patches. These patterns support the existence of scale-dependent feedbacks in Chihuahuan Desert grasslands and indicate an optimal patch size for reproductive effort and success in B. eriopoda . We discuss the implications of these results for detecting ecological thresholds in desert grasslands.

New Mexico

Forest reproduction along a climatic gradient in the Sierra Nevada, California

To elucidate broad-scale environmental controls of coniferous forest reproduction in the Sierra Nevada, California, we monitored reproduction for 5 years in 47 plots arrayed across a steep elevational (climatic) gradient. We found that both absolute seedling densities (stems < 1.37 m) and seedling densities relative to overstory parent tree basal area declined sharply with elevation. Rates of seedling turnover (the average of birth and death rates) also declined with elevation. In contrast, seed production was not predicted by elevation and was highly variable from year to year. During a mast year of seed production, the intensity of masting was uneven among plots. Seedling densities were elevated only during the single year immediately following the mast year, suggesting reproduction in our forests may be primarily limited by abiotic factors such as the availability of suitable sites and weather. Disturbance also clearly affected reproduction; plots that had recently burned had significantly higher seedling to parent tree ratios for Abies species, suggesting that even though established Abies concolor may be relatively susceptible to fire, the species can recover rapidly through prolific reproduction. Since reproductive failures may be our earliest signal of changing forest conditions, seedling dynamics could provide a sensitive, if variable, indicator of environmental changes.

California

Stress and fish reproduction: The roles of allostasis and hormesis

This paper is a review of the effects of stress on reproduction in fishes. I hope to further the development of the concepts of allostasis and hormesis as relevant to understanding reproduction in general and in fish in particular. The main contentions I derive in this review are the following: Stressors affect fish reproduction in a variety of ways depending on the nature and severity of the stressor. The effects are transduced through a hormonal cascade initiated by perception of the stressor and involving the hypothalamus-pituitary-interrenal axis, the catecholamines, and also cytokines. Mounting a stress response and resisting a stressor is an energetically costly process, including costs associated with allostasis, attempting to reset homeostatic norms. Responses in emergency situations (e.g., being chased by a predator or a net) can be different from those where fish can cope (e.g., being in a more crowded environment) with a stressor, but both situations involve energy re-budgeting. Emergency responses happen in concert with the onset of energy limitations (e.g., the fish may not eat), while coping with allostatic overload can happen in a more energy-rich environment (e.g., the fish can continue to eat). Low levels of stress may have a positive effect on reproductive processes while greater stress has negative effects on fish reproduction. The concept of hormesis is a useful way to think about the effect of stressors on fish reproduction since responses can be nonmonotonal, often biphasic.

General and Comparative Endocrinology

Aqueous exposure to the progestin, levonorgestrel, alters anal fin development and reproductive behavior in the eastern mosquitofish (Gambusia holbrooki)

Endogenous progestogens are important regulators of vertebrate reproduction. Synthetic progestins are components of human contraceptive and hormone replacement pharmaceuticals. Both progestogens and progestins enter the environment through a number of sources, and have been shown to cause profound effects on reproductive health in various aquatic vertebrates. Progestins are designed to bind human progesterone receptors, but they also have been shown to strongly activate androgen receptors in fish. Levonorgestrel (LNG) activates fish androgen receptors and induces development of male secondary sex characteristics in females of other species. Although behavior has been postulated to be a sensitive early indicator of exposure to certain environmental contaminants, no such research on the reproductive behavior of gestagen-exposed fish has been conducted to date. The goal of our study was to examine the exposure effects of a human contraceptive progestin, LNG, on the reproductive development and behavior of the viviparous eastern mosquitofish ( Gambusia holbrooki ). Internal fertilization is a requisite characteristic of viviparous species, and is enabled by an androgen driven elongation of the anal fin into the male gonopodium (i.e., phallus). In this study, we exposed adult mosquitofish to ethanol (EtOH control), 10 ng/L, and 100 ng/L LNG for 8 d using a static replacement exposure design. After 8 d, a subset of males and females from each treatment were examined for differences in the 4:6 anal fin ratio. In addition, paired social interaction trials were performed using individual control males and control females or females treated 10 ng/L or 100 ng/L LNG. Female mosquitofish exposed to LNG were masculinized as evidenced by the elongation of the anal fin rays, a feature normal to males and abnormal to females. LNG caused significant increases in the 4:6 anal fin ratios of female mosquitofish in both the 10 ng/L and 100 ng/L treatments, although these differences were not significant between the two treatments. LNG caused significant increases in the 4:6 anal fin ratio of males exposed to 100 ng/L, with no effects observed in the 10 ng/L treatment. In addition, the reproductive behavior of control males paired with female mosquitofish exposed to 100 ng/L LNG was also altered, for these males spent more time exhibiting no reproductive behavior, had decreased attending behavior, and a lower number of gonopodial thrusts compared to control males paired to control female mosquitofish. Given the rapid effects on both anal fin morphology and behavior observed in this study, the mosquitofish is an excellent sentinel species for the detection of exposure to LNG and likely other 19-nortestosterone derived contraceptive progestins in the environment.

General and Comparative Endocrinology

An evaluation of biomarkers of reproductive function and potential contaminant effects in Florida largemouth bass (Micropterus salmoides floridanus) sampled from the St. Johns River

The objective of this study was to describe and compare several reproductive parameters for Florida largemouth bass ( Micropterus salmoides floridanus ) inhabiting the St. Johns River and exposed to different types and/or degrees of contamination. Welaka was selected as the reference site in this study because of its low urban and agricultural development, Palatka is in close proximity to a paper mill plant, the Green Cove site is influenced by marine shipping activities and Julington Creek site receives discharges of domestic wastewater and storm water runoff from recreational boating marinas. For this study, bass were sampled both prior to (September 1996) and during the spawning season (February 1997). In order to characterize chemical exposure, bass livers were analyzed for up to 90 trace organics and 11 trace metal contaminants. Reproductive parameters measured included gonadosomatic index (GSI), histological evaluation of gonads and plasma concentrations of vitellogenin (VTG), 17β-estradiol (E 2 ) and 11-ketotestosterone (11-KT). In general, the sum of organic chemicals was highest in livers from Palatka bass and bass from Green Cove and Julington Creek had higher hepatic concentrations of low molecular polycyclic aromatic hydrocarbons and polychlorinated biphenyls when compared to fish from Welaka. Metals were more variable across sites, with highest mean concentrations found in bass from either Julington Creek (Ag, As, Cr, Cu, Zn) or Welaka (Cd, Hg, Pb, Se, Tn). Female bass from Palatka and Green Cove had lower concentrations of E 2 , VTG and lower GSI in relation to Welaka. Males from Palatka and Green Cove showed comparable declines in 11-KT in relation to males from Julington Creek and GSI were decreased only in Palatka males. These results indicate a geographical trend in reproductive effects, with changes being most pronounced at the site closest to the paper mill (Palatka) and decreasing as the St. Johns River flows downstream. Since reproductive alterations were most evident in bass sampled from the site closest to the paper mill discharge, it is possible that exposure to these effluents might explain at least some of the results reported here. However, the presence of reproductive alterations in fish sampled at a considerable distance from the mill discharge (Green Cove, 40 km) would suggest exposure to chemicals released from sources other than the paper mill plant. It is clear that additional studies are needed to evaluate the potential impact of these reproductive changes in populations of Florida largemouth bass inhabiting the St. Johns River.

Florida

Reproductive allochrony in seasonally sympatric populations maintained by differential response to photoperiod: Implications for population divergence and response to climate change

Reproductive allochrony presents a potential barrier to gene flow and is common in seasonally sympatric migratory and sedentary birds. Mechanisms mediating reproductive allochrony can influence population divergence and the capacity of populations to respond to environmental change. We asked whether reproductive allochrony in seasonally sympatric birds results from a difference in response to supplementary or photoperiodic cues and whether the response varies in relation to the distance separating breeding and wintering locations as measured by stable isotopes. We held seasonally sympatric migratory and sedentary male dark-eyed juncos ( Junco hyemalis ) in a common garden in early spring under simulated natural changes in photoperiod and made measurements of reproductive and migratory physiology. On the same dates and photoperiods, sedentary juncos had higher testosterone (initial and gonadotropin-releasing hormone induced), more developed cloacal protuberances, and larger testes than migrants. In contrast, migratory juncos had larger fat reserves (fuel for migration). We found a negative relationship between testis mass and feather hydrogen isotope ratios, indicating that testis growth was more delayed in migrants making longer migrations. We conclude that reproductive allochrony in seasonally sympatric migratory and sedentary birds can result from a differential response to photoperiodic cues in a common garden, and as a result, gene flow between migrants and residents may be reduced by photoperiodic control of reproductive development. Further, earlier breeding in response to future climate change may currently be constrained by differential response to photoperiodic cues.

American Naturalist

Variations in the reproductive cycle of Dreissena polymorpha in Europe, Russia, and North America

The reproductive cycle of the zebra mussel { Dreissena polymorpha ) is highly variable throughout its range in Europe, Russia , and North America. The environmental factors influencing this variation are poorly understood, but successful reproduction is occurring in areas where it was initially believed that adult zebra mussels could not survive (i.e., southern United States). The differences in mussel reproduction occurring from site-to-site make it difficult to predict timing of specific events, such as the start of larval production, that are important in initiating containment or control procedures. For example, the amount of time required for a fertilized egg to develop into a juvenile mussel can be as short as 8 days, or as long as 240 days. Release of gametes by adults can be a highly synchronized event, focused over a 1–2 week period, or it can be completely non-synchronized, occurring throughout the year. Zebra mussels in some localities start spawning at water temperatures of 12–13°C, but do not start until water temperatures reaches 22°C at other sites. While some of this variability in reproductive behavior stems from mussel adaptation to local conditions, part is due to difficulties in sampling these events. It is difficult to determine reproductive success of a specific population because of the problems in separating locally produced larvae from larvae drifting in from other areas. Further research is needed not only on the relationship between reproduction and environment at the community level, but also on the variability in response of individual mussels.

American Zoologist

Reproduction in moose at their southern range limit

Reproduction is a critical fitness component in large herbivores. Biogeographic models predict that populations occurring at the edges of the range may have compromised reproductive rates because of inferior habitat at range peripheries. When reproductive rates are chronically low, ungulate populations may lack the resiliency to rebound quickly after periods of environmental stress, and this effect may be greatest for heat-sensitive organisms at their southern range limit. To assess the demographic vulnerability of moose ( Alces alces ), we studied relationships between reproductive rates, maternal age, and rump fat in the southernmost naturally occurring moose population in North America. For prime-aged moose in our study, pregnancy rates were high (92%), but moose aged < 3 or > 9 years had low pregnancy rates (32% and 38%, respectively). The relationship between rump fat and pregnancy was nonlinear such that a threshold of at least 2mm of rump fat yielded a high probability of being pregnant midwinter. In contrast, among pregnant moose, the probability of both producing a calf and recruiting it until spring increased linearly with rump fat. We also conducted a meta-analysis of pregnancy and twinning rates for adult (≥ 2 years) moose across a latitudinal gradient to compare reproductive rates from our study to other populations in North America. Moose living at southern latitudes tended to have lower reproductive rates than those living in the core of moose range, implying that southern moose populations may be demographically more vulnerable than northern moose populations.

Utah