Effects of reproduction on the antipredator tactics of snakes (Tropidonophis mairii, Colubridae)

Does an animal’s reproductive state influence the distance at which it flees from an approaching predator? We predicted that reproduction would increase approach distances in pregnant females (because they are burdened with eggs and thus less able to escape rapidly), but reduce them in males (because of lower vigilance due to males focusing on mate-searching rather than predator detection). Field data on approach distances of keelback snakes (Tropidonophis mairii, Natricinae, Colubridae) supported both of these predictions. We walked the same 1.3-km transect along a dam wall in tropical Australia on 135 nights, and recorded the distances at which snakes fled from our approach. Locomotor speeds were measured for a subset of these animals. Variations in approach distance due to season, weather conditions, prior capture history and snake body size were minor, but reproduction strongly affected approach distances for snakes of both sexes. Gravid females were slower than non-gravid females, and fled at greater distances. Reproductive status did not affect locomotor speeds of males, but males that were reproductive (i.e., contained sperm) permitted closer approach than did non-reproductive adult males. Reproduction thus affected approach distances in snakes of both sexes, but in opposite directions and for different reasons.     

Estimating survival rates of uncatchable animals: the myth of high juvenile mortality in reptiles

Survival rates of juvenile reptiles are critical population parameters but are difficult to obtain through mark-recapture programs because these small, secretive animals are rarely caught. This scarcity has encouraged speculation that survival rates of juveniles are very low, and we test this prediction by estimating juvenile survival rates indirectly. A simple mathematical model calculates the annual juvenile survival rate needed to maintain a stable population size, using published data on adult survival rates, reproductive output, and ages at maturity in 109 reptile populations encompassing 57 species. Counter to prediction, estimated juvenile survival rates were relatively high (on average, only about 13% less than those of conspecific adults) and highly correlated with adult survival rates. Overall, survival rates during both juvenile and adult life were higher in turtles than in snakes, and higher in snakes than in lizards. As predicted from life history theory, rates of juvenile survival were higher in species that produce large offspring, and higher in viviparous squamates than in oviparous species. Our analyses challenge the widely held belief that juvenile reptiles have low rates of annual survival and suggest instead that sampling problems and the elusive biology of juvenile reptiles have misled researchers in this respect.     

Variation in size at sex-change among natural populations of the protandrous hermaphrodite, <Emphasis Type="Italic">Crepidula fornicata</Emphasis> (Gastropoda,Calyptraeidae)

Protandrous hermaphrodites are predicted to change sex from male to female when relative reproductive fitness of females surpasses that of males. How size at sex transition varies with population, mating group and individual parameters was investigated for five populations of the protandrous hermaphrodite slipper snail, Crepidula fornicata. The populations varied for density, size distribution, average mating group size and sex ratio. Size at sex-change was correlated with the population sex ratio. Comparisons of multiple hypotheses revealed that variables predicting the sex of a snail vary among positions in the mating group. The variables included body size, the relative size of the snail sitting atop the focal snail and population density. Our data support the conclusions that size at sex-change (and by inference, the size at which one sex has relatively greater fitness) is not fixed for these hermaphrodites and that individual size, social conditions and population differences all influence variation in relative fitness.     

Life-history adaptations to arboreality in snakes

If selective forces on locomotor ability and reproductive biology differ among habitats, we expect to see relationships between habitat, morphology, and life-history traits. Comparative (phylogenetically based) analysis of data from 12 pythonid and 12 boid snake species reveals multiple evolutionary shifts in habitat use, notably in the evolution of arboreal habits. Compared to terrestrial and aquatic taxa of the same overall body size, arboreal species have narrower and more laterally compressed bodies and relatively longer tails. Offspring sizes are not affected by arboreality, but presumably reflecting space constraints within their narrow bodies, arboreal species (1) produce smaller clutch sizes relative to maternal body length and (2) have left and right ovaries that overlap little if at all along the length of the body (i.e., the right ovary is positioned anterior to the left ovary) whereas in terrestrial snakes the two ovaries overlap along much of their length. This modification of ovarian morphology in arboreal snakes presumably reduces the degree of bodily distension during vitellogenesis and pregnancy, thus enhancing climbing ability and camouflage among the branches.     

Dominance rank,female reproductive synchrony,and male reproductive skew in wild Assamese macaques

In groups with multiple males, direct mate competition may select for the evolution of dominance hierarchies that sort males into a queue for access to fertile females. The priority-of-access (PoA) model proposed by Altmann in 1962 makes explicit predictions about the resulting paternity distribution based on an interaction between male dominance rank and the overlap of female receptive phases. Here, we investigated whether the logic of the PoA model predicted the distribution of paternity across ranks in a seasonal breeder with high reproductive synchrony over six consecutive mating seasons. We studied 18 males that resided in a group of wild Assamese macaques (Macaca assamensis) in their natural habitat at Phu Khieo Wildlife Sanctuary, Thailand, between 2006 and 2011 with 5 to 13 conceptions per season. We assessed whether mate guarding increased paternity success, described “short-term” deviations from predicted paternity distribution, and examined how these are related to the number of competitors and fertile females. We determined genetic paternity of 43 (93 %) offspring born into the study group and found reproductive skew to be relatively low with 29 % alpha male paternity in accordance with the high degree of female reproductive synchrony observed. Short-term deviations from expected paternity distribution over ranks were not explained by the number of resident males or the number of conceiving females or their interaction. Within the limits of this study, these results suggest that even if males cannot discern female fertile phases, if reproduction is seasonal, and if reproductive synchrony is high, males may also compete directly over access to females.     

Esterase heterogeneity and an investigation into racial differences in the cirripede Balanus balanoides using acrylamide gel electrophoresis

Four groups of bands showing esterase activity were found after electrophoretic separation of extracts of Balanus balanoides (L.) prosoma on polyacrylamide gels. Each group was distinguishable from other groups by mobility, reaction with specific substrates, and susceptibility to various inhibitors. Groups designated BbEII and BbEIII showed cholinesterase activity, BbEIV showed carboxylesterase activity, and BbEI possible arylesterase activity. Polymorphisms were found within Groups BbEIII and BbEI. The frequency distribution of BbEIV polymorphs in a sample from the Menai Straits, UK, fitted the Hardy-Weinberg predicted values for 4 co-dominant alleles. A sample from St. Andrews, New Brunswick, Canada, however, did not fit the Hardy-Weinberg prediction. No variation between individuals was found when malate dehydrogenase (MDH) and lactate dehydrogenase (LDH) isozymes were examined. Hence, these systems are unsuitable for testing racial differences. The results for the cholinesterase and arylesterase isozymes substantiate the view, originally based on differences in reproductive phenology and egg size, that genetically separate races exist on either side of the Atlantic Ocean.     

A structural equation model to integrate changes in functional strategies during old-field succession

From a functional perspective, changes in abundance, and ultimately species replacement, during succession are a consequence of integrated suites of traits conferring different relative ecological advantages as the environment changes over time. Here we use structural equations to model the interspecific relationships between these integrated functional traits using 34 herbaceous species from a Mediterranean old-field succession and thus quantify the notion of a plant strategy. We measured plant traits related to plant vegetative and reproductive size, leaf functioning, reproductive phenology, seed mass, and production on 15 individuals per species monitored during one growing season. The resulting structural equation model successfully accounts for the pattern of trait covariation during the first 45 years post-abandonment using just two forcing variables: time since site abandonment and seed mass; no association between time since field abandonment and seed mass was observed over these herbaceous stages of secondary succession. All other predicted traits values are determined by these two variables and the cause-effect linkage between them. Adding pre-reproductive vegetative mass as a third forcing variable noticeably increased the predictive power of the model. Increasing the time after abandonment favors species with increasing life span and pre-reproductive biomass and decreasing specific leaf area. Allometric coefficients relating vegetative and reproductive components of plant size were in accordance with allometry theory. The model confirmed the trade-off between seed mass and seed number. Maximum plant height and seed mass were major determinants of reproductive phenology. Our results show that beyond verbal conceptualization, plant ecological strategies can be quantified and modeled.     

Species abundance distributions result from body size-energetics relationships

Abundance distributions are a central characteristic of ecosystems. Certain distributions have been derived from theoretical models of community organization, and therefore the fit of data to these distributions has been proposed as a test of these theories. However, it is shown here that the geometric sequence distribution can be derived directly from the empirical relationship between population density and body size, with the assumption of random or uniform body size distributions on a log scale (as holds at local scales). The geometric sequence model provides a good to excellent fit to empirical data. The presence of noise in the relationship between population density and body size creates a curve that begins to approximate a lognormal species abundance distribution as the noise term increases. For continental-scale data in which the body size distribution is not flat, the result of sampling tends again toward the lognormal. Repeat sampling over time smooths out species population fluctuations and damps out the noise, giving a more precise geometric sequence abundance distribution. It is argued that the direct derivation of this distribution from empirical relationships gives it priority over distributions derived from complex theoretical community models.     

Ideal dominance distributions: a test using red-winged blackbirds (Agelaius phoeniceus)

Summary The ideal dominance distribution model predicts that competition between individuals of a species for territories will result in socially dominant individuals acquiring territories in higher quality habitat than their subordinates. Although the dispersion and relative reproductive success of male red-winged blackbirds (Agelaius phoeniceus) across habitats in eastern Ontario appears to conform to the ideal dominance distribution model, data from a study of three captive groups comprised of males from both high (marsh) and low (upland field) quality habitats failed to support the prediction that males from marsh habitat are dominant to those from upland habitat. Contrary to the prediction males from uplands were generally dominant to males from marshes. We found a significant positive correlation between dominance and both increased epaulet size and increased body size. Controlling for these positive effects, upland males remained generally dominant to marsh males. Measurements of independent samples of males from both habitats indicated that the overall distribution of males does not conform to ideal dominance. We suggest that the strong between-year territory fidelity shown by male red-winged blackbirds and chance events when they initially acquire territories may contribute to this lack of conformity.     

Different modes of resource variation provide a critical test of ideal free distribution models

Ideal free distribution (IFD) models are perhaps the group of mathematical models of behavior that have been the most widely and successfully applied by empiricists. These models can be applied to nearly any situation in which consumers compete—by any mechanism—for resources that are patchily distributed in their environment. Although IFD models have come to be broadly accepted, experiments that simultaneously test more than a single prediction are rare. Instead, investigators normally either test (1) for a relationship between the distribution of consumers and the distribution of resources or (2) whether average fitnesses are equal across resource patches. We conducted experiments with pea aphids (Acyrthosiphon pisum Harris) feeding on two patches of fava beans (Vicia faba L.) to fully independently parameterize an IFD model with interference competition and then test quantitative predictions about aphid spatial distributions and the payoffs of patch choice. We found a precise fit between aphids’ predicted and observed reproductive successes. Furthermore, by varying patch “quality” in two ways, we were able to show that aphid distributions vary with the mode of resource variation in the predicted manner: aphids (1) matched resources when patches varied in size but not quality and (2) overmatched the good patch when patches varied in quality but not size (predicted as a consequence of weak interference). The close correspondence between quantitative predictions of the model with observed behaviors suggests that IFD theory is a framework with more explanatory power than is generally appreciated.     

Prey morphology constrains the feeding ecology of an aquatic generalist predator

Resource availability and accessibility are primary factors guiding the distribution and abundance of organisms. For generalists, prey availability reflects both prey abundance and differences in quality among prey taxa. Although some aspects of prey quality, such as nutritional composition, are well studied, our understanding of how prey morphology contributes to overall prey quality is limited. Because snakes cannot reduce prey size by mastication, many aspects of their feeding ecology (e.g., maximum prey size, feeding performance, and the degree of postprandial locomotor impairment) may be affected by prey shape. We conducted a uniquely comprehensive comparison of prey quality for a generalist species, the banded watersnake (Nerodia fasciata), using prey that were similar in mass and presumably similar in nutritional composition but different in shape and habitat association. Specifically, we compared nutritional composition and shape of paedomorphic salamanders (Ambystoma talpoideum) and sunfish (Lepomis MARGINATUS) and used a series of repeated-measures experiments to examine feeding performance (number of prey consumed, maximum prey size, and intra-oral transport time), digestive metabolism (specific dynamic action, SDA), and postprandial locomotor performance of snakes fed Ambystoma and Lepomis. Cost of digestion was similar between the prey types, likely reflecting their similar nutritional composition. However, snakes consumed larger Ambystoma than Lepomis and intra-oral transport time was much shorter for Ambystoma. Snakes fed Lepomis also suffered greater reduction in crawling speed than those fed Ambystoma. These differences highlight the need for behaviorally integrated approaches to understanding prey quality and support field observations of the importance of amphibian prey for juvenile watersnakes.     

Migration effects in a stochastic multipopulation model for African bee population dynamics

The rate of northern migration of the Africanized honey bee (AHB) in the United States has recently slowed dramatically. This paper investigates the impact of migration on the equilibrium size distributions of a particular stochastic multipopulation model, namely a coupled logistic power law model. The bivariate equilibrium size distribution of the model is derived and illustrated with parameter values used to describe AHB population dynamics. In the model, the difference between the equilibrium sizes of the two populations is a measure of the effect of migration. The distribution of this difference may be approximated by a normal distribution. The mean and variance parameters for the normal are predicted accurately by a second-order regression model based on the migration rate and the maximum size of the first population. The methodology is general, and should be useful in studying the migration effect in many other applications with one-way migration.     

Natural variation in the concentrations of haemocyanin from three decapod crustaceans, Nephrops norvegicus, Liocarcinus depurator and Hyas aranaeus

Natural variation in the concentrations of haemocyanin ([Hc]) is examined for three decapod species collected from two different locations (Gullmarsfjord and Kattegat) off the west coast of Sweden (August to September 1998). Only one of the frequency distributions for [Hc] is normally distributed, and median values differ both between and within species. Differences in [Hc] cannot be attributed to sex, reproductive condition or moulting condition (over the limited range of moult stages examined) for any of the species. While body size did not scale with [Hc] for the Norway lobster Nephrops norvegicus or the swimming crab Liocarcinus depurator, it did for the spider crab Hyas araneus. To our knowledge this is the first time a negative relationship between body size and [Hc] has been reported for any crustacean. Thus the right-skewed frequency distribution of spider crab [Hc] could be accounted for by the right-skewed body size distribution recorded. The shape of the frequency distributions for [Hc] from the other two species could not be accounted for through differences in the factors examined here, although it is suggested that the amount of food consumed (or not) may be important. Received: 19 July 1999 / Accepted: 4 October 1999     

Polyandry and paternity in a wild population of the swordtail Xiphophorus nigrensis

Genetic analyses of parentage provide crucial information about the prevalence of polyandry and the potential for sexual selection to operate in wild populations. In the swordtail Xiphophorus nigrensis, large males are thought to have a competitive advantage due to their superiority in male–male contests and attractiveness to females, who are presumed to mate multiply. I examined the distribution of paternity within broods, the relationship between male body size and paternity and the effect of sire number on fecundity from females collected in the field. Sixty-one percent of females produced offspring from two to four males, with 70% of the offspring typically sired by one of the males represented in the brood. Male body size did not affect paternity share or whether females were multiply mated, as predicted if precopulatory sexual selection has a strong effect on the outcome of postcopulatory sexual selection. Female fecundity increased with the number of sires; however, this relationship was not observed when the smallest broods, where multiple mating is more difficult to detect, were excluded from the analysis. The high levels of multiple paternity and reproductive skew suggest that postcopulatory sexual selection has important evolutionary consequences in X. nigrensis. Traits important in precopulatory sexual selection, such as male body size, however, are more likely to affect sexual selection by increasing the number of mates obtained rather than paternity share within broods.     

Patterns of Diversity of Neotropical Squamate Reptile Species with Emphasis on the Brazilian Amazon and the Conservation Potential of Indigenous Reserves

We present new distibutional data for squamate reptiles (amphisbaenians, lizards, and snakes) from several Neotropical sites and estimate species (alpha) diversity at each site. Species compositions at all localities were used to estimate similarity in species richness within and between each of the major regions sampled: Central American lowland forests, Amazonia, and South American dryland forests. We used these estimates to derive maps summarizing overall regional patterns (beta diversity) of squamate richness. Both groups clearly reveal distinct faunal divisions among the three regions. Within Amazonia, lizards and snakes show a clear division in species composition between a "western edge" cluster of sites adjacent to eastern Andean slopes and all other localities in the central and eastern regions. Within this east-central block of sites, a further division is evident between localities north and south of the main Amazon River channel. These general regions of squamate similarity were qualitatively assessed in Brazil's portion of Amazonia within the context of the distribution of all state, federal, and Indian lands that are in principle afforded some level of protection and therefore offer potential for biodiversity conservation. We assess the potential for biodiversity conservation relative to the size of reserves needed to sustain populations of 500 individuals of the three largest species of snakes and to the distribution of potential conservation areas relative to the regional patterns of diversity identified by the statistical analysis. We conclude that the sizes and distributions of reserves are adequate to sustain viable populations of squamates if these areas can be counted on to play a major role in biodiversity conservation.     

Sperm allocation in relation to male traits, female size, and copulation behaviour in freshwater crayfish species

Sperm competition is a well-recognised agent in the evolution of sperm and ejaculate structure, as well as variation in female quality. Models of the evolution of ejaculate expenditure predict that male body condition, female fecundity and the risk and intensity of sperm competition may be the ultimate factors shaping optimal ejaculate size. We investigated sperm allocation in Austropotamobius italicus, a freshwater crayfish exhibiting a coercive mating system and external fertilisation, in relation to male and female traits and copulation behaviour under laboratory conditions. We found that mating males were sensitive to female size and produced larger ejaculates when mating with larger females, which were more fecund in terms of number of eggs produced. We found no evidence for female egg production being sperm-limited, as the number of eggs was not dependent on male sperm expenditure. Copulation duration and number of ejaculations reliably predicted the amount of sperm transferred, and both these behavioural measures positively covaried with female body size. These results indicate that male freshwater crayfish can modulate their sperm expenditure in accordance with cues that indicate female fecundity. In addition, a novel finding that emerged from this study is the decrease in sperm expenditure with male body size, which may either suggest that large, old male crayfish are better able than small males to economise sperm at a given mating to perform multiple matings during a reproductive season, or that they experience senescence of their reproductive performance.     

Do predators control prey species abundance? An experimental test with brown treesnakes on Guam

The effect of predators on the abundance of prey species is a topic of ongoing debate in ecology; the effect of snake predators on their prey has been less debated, as there exists a general consensus that snakes do not negatively influence the abundance of their prey. However, this viewpoint has not been adequately tested. We quantified the effect of brown treesnake (Boiga irregularis) predation on the abundance and size of lizards on Guam by contrasting lizards in two 1-ha treatment plots of secondary forest from which snakes had been removed and excluded vs. two 1-ha control plots in which snakes were monitored but not removed or excluded. We removed resident snakes from the treatment plots with snake traps and hand capture, and snake immigration into these plots was precluded by electrified snake barriers. Lizards were sampled in all plots quarterly for a year following snake elimination in the treatment plots. Following the completion of this experiment, we used total removal sampling to census lizards on a 100-m2 subsample of each plot. Results of systematic lizard population monitoring before and after snake removal suggest that the abundance of the skink, Carlia ailanpalai, increased substantially and the abundance of two species of gekkonids, Lepidodactylus lugubris and Hemidactylus frenatus, also increased on snake-free plots. No treatment effect was observed for the skink Emoia caeruleocauda. Mean snout-vent length of all lizard species only increased following snake removal in the treatment plots. The general increase in prey density and mean size was unexpected in light of the literature consensus that snakes do not control the abundance of their prey species. Our findings show that, at least where alternate predators are lacking, snakes may indeed affect prey populations.     

Partitioning the factors of spatial variation in regeneration density of shade-tolerant tree species

Understanding coexistence of highly shade-tolerant tree species is a longstanding challenge for forest ecologists. A conceptual model for the coexistence of sugar maple (Acer saccharum) and American beech (Fagus grandibfolia) has been proposed, based on a low-light survival/high-light growth trade-off, which interacts with soil fertility and small-scale spatiotemporal variation in the environment. In this study, we first tested whether the spatial distribution of seedlings and saplings can be predicted by the spatiotemporal variability of light availability and soil fertility, and second, the manner in which the process of environmental filtering changes with regeneration size. We evaluate the support for this hypothesis relative to the one for a neutral model, i.e., for seed rain density predicted from the distribution of adult trees. To do so, we performed intensive sampling over 86 quadrats (5 x 5 m) in a 0.24-ha plot in a mature maple-beech community in Quebec, Canada. Maple and beech abundance, soil characteristics, light availability, and growth history (used as a proxy for spatiotemporal variation in light availability) were finely measured to model variation in sapling composition across different size classes. Results indicate that the variables selected to model species distribution do effectively change with size, but not as predicted by the conceptual model. Our results show that variability in the environment is not sufficient to differentiate these species' distributions in space. Although species differ in their spatial distribution in the small size classes, they tend to correlate at the larger size class in which recruitment occurs. Overall, the results are not supportive of a model of coexistence based on small-scale variations in the environment. We propose that, at the scale of a local stand, the lack of fit of the model could result from the high similarity of species in the range of environmental conditions encountered, and we suggest that coexistence would be stable only at larger spatial scales at which variability in the environment is greater.     

Selective harvesting and habitat loss produce long-term life history changes in a mouflon population.

We examined the long-term effects (28 years) of habitat loss and phenotype-based selective harvest on body mass, horn size, and horn shape of mouflon (Ovis gmelini musimon) in southern France. This population has experienced habitat deterioration (loss of 50.8% of open area) since its introduction in 1956 and unrestricted selective hunting of the largest horned males since 1973. Both processes are predicted to lead to a decrease in phenotype quality by decreasing habitat quality and by reducing the reproductive contribution of individuals carrying traits that are targeted by hunters. Body mass and body size of both sexes and horn measurements of males markedly decreased (by 3.4-38.3%) in all age classes from the 1970s. Lamb body mass varied in relation to the spatiotemporal variation of habitat closure within the hunting-free reserve, suggesting that habitat closure explains part of these changes. However, the fact that there was no significant spatial variation in body mass in the early part of the study, when a decline in phenotypic quality already had occurred, provided support for the influence of selective harvesting. We also found that the allometric relationship between horn breadth and horn length changed over the study period. For a given horn length, horn breadth was lower during the second part of the study. This result, as well as changes in horn curve diameter, supports the interpretation that selective harvesting of males based on their horn configuration had evolutionary consequences for horn shape, since this phenotypic trait is less likely to be affected by changes in habitat characteristics. Moreover, males required more time (approximately four years) to develop a desirable trophy, suggesting that trophy hunting favors the reproductive contribution of animals with slow-growing horns. Managers should exploit hunters' desire for trophy males to finance management strategies which ensure a balance between the population and its environment. However, for long-term sustainable exploitation, harvest strategy should also ensure that selectively targeted males are allowed to contribute genetically to the next generations.     

Body condition and the grouping behavior of brood-caring female common eiders (<Emphasis Type="Italic">Somateria mollissima</Emphasis>)

Both theoretical and empirical work has shown that group size increases with increasing ecological constraints on solitary breeding. Ecological constraints refer to extrinsic factors such as availability of breeding sites, food or mates. Common eider (Somateria mollissima) females pool their broods and share brood-rearing duties, or rear broods alone. Females are often in poor condition at hatching, as incubation is accomplished without feeding, and variation in body condition is largely environmentally induced and thus unpredictable. We found that the intensity of and duration of parental care that females provide is positively correlated with their body condition at hatching. This suggests that body condition is an ecological constraint on successful solitary breeding. We further observed that group productivity in common eider broods is a decelerating function of the number of tending females. As predicted, females in poorer condition (i.e., facing stronger ecological constraints) were found in larger groups. This result is straightforward if solitary tenders can enter any group at no cost. However, if entry is group-controlled, stable groups of non-relatives are predicted not to occur when per capita reproduction declines with group size. The N-person staying incentive model permits groups to form under these conditions, because reproduction is unevenly divided between dominants and subordinates in the group. We discuss the plausibility of these alternative models of group size for understanding the grouping behavior of brood-caring female common eiders.Communicated by M. Webster