Reproductive consequences of variation in the composition of howler monkey (Alouatta spp.) groups

At least three general categories of environmental pressure - predation, resource distribution, and demographics - shape the costs and benefits of group-living for animals. Among the demographic factors that influence individual survival and reproduction, the composition of social groups can play an important role. Census data drawn from 26 populations of howler monkeys (Alouatta spp.) were used to determine if the composition of groups explained variation in their reproductive performance. Each group's reproductive performance was estimated by calculating the difference between the observed number of immatures and the number expected from its population average. Of four group structure variables tested, only one - the residual of the adult and subadult sex ratio - was a consistent correlate of reproductive performance across the howler monkey populations. Groups with a greater proportion of adult and subadult males contained more juveniles than expected from the population average. I propose that the survival or retention of immatures within howler monkey groups depends in part on the behavior of resident males. Of particular importance, the relative proportions of resident males and females were more informative than the absolute number of males or females. On this basis, I evaluate the possible role of males in protection from predation, conspecific aggression, and resource competition. The techniques used here can also be used to forecast major changes in demographic structure within populations.     

Male backspace availability in the giant waterbug (Belostoma flumineum Say)

Summary Following bouts of courtship and copulation, female giant waterbugs (Belostoma flumineum Say) deposit eggs on the backs of their mates. Throughout a 6– to 12-day brooding period, males display several behaviors that are vital to egg-nymph survival. Consequently, females depend on male post-copulatory behaviors for successful reproduction and the possibility exists for male backspace availability to limit female reproduction in this species. I studied seasonal trends and factors that affect male backspace availability in populations of B. flumineum in east-central Illinois (USA). Early in the spring/summer, giant waterbug populations are relatively small and a large majority (188/205=91.7%) of the males are egg-laden; males experimentally added to the population during this period quickly became encumbered. In contrast, later in the summer after young-of-the-year emerge as adults, the waterbug population density increases dramatically and fewer (670/1274\2= 52.6%) of the males are encumbered (egg-laden). Of the males that are egg-laden both early and late in the season, significantly more are completely encumbered (i.e., 100% of the dorsum egg covered) early in the spring. The adult sex ratio is generally not biased and the number of eggs/pad that completely covers a male approximates a full ovarian complement. Therefore, these factors probably do not cause male backspace to become limited. The primary factor that appears to limit male backspace availability is the ability of females to synthesize a second partial clutch in a short time, often within 1 to 4 days. Females are capable of ovipositing partial clutches on 12 males within 30 days, whereas male brooding period is temperature dependent and ranges from 6 to 13 days. Newly emerged males are capable of breeding significantly sooner than can newly emerged females, thereby creating ample oviposition substrate for the females in the population after young-of-the-year adults appear. The evolution of sex-role reversal is not well understood; however it should not evolve in waterbugs unless male backspace limits female reproduction. Such a situation appears to exist in B. flumineum early in the season but not later in the summer.     

Effects of Multiple Levels of Social Organization on Survival and Abundance

Abstract: Identifying how social organization shapes individual behavior, survival, and fecundity of animals that live in groups can inform conservation efforts and improve forecasts of population abundance, even when the mechanism responsible for group‐level differences is unknown. We constructed a hierarchical Bayesian model to quantify the relative variability in survival rates among different levels of social organization (matrilines and pods) of an endangered population of killer whales (Orcinus orca). Individual killer whales often participate in group activities such as prey sharing and cooperative hunting. The estimated age‐specific survival probabilities and survivorship curves differed considerably among pods and to a lesser extent among matrilines (within pods). Across all pods, males had lower life expectancy than females. Differences in survival between pods may be caused by a combination of factors that vary across the population's range, including reduced prey availability, contaminants in prey, and human activity. Our modeling approach could be applied to demographic rates for other species and for parameters other than survival, including reproduction, prey selection, movement, and detection probabilities.     

Effects of age and sex ratios on offspring recruitment rates in translocated black rhinoceros

Success of animal translocations depends on improving postrelease demographic rates toward establishment and subsequent growth of released populations. Short‐term metrics for evaluating translocation success and its drivers, like postrelease survival and fecundity, are unlikely to represent longer‐term outcomes. We used information theory to investigate 25 years of data on black rhinoceros (Diceros bicornis) translocations. We used the offspring recruitment rate (ORR) of translocated females—a metric integrating survival, fecundity, and offspring recruitment at sexual maturity—to detect determinants of success. Our unambiguously best model (AICω = 0.986) predicted that ORR increases with female age at release as a function of lower postrelease adult rhinoceros sex ratio (males:females). Delay of first postrelease reproduction and failure of some females to recruit any calves to sexual maturity most influenced the pattern of ORRs, and the leading causes of recruitment failure were postrelease female death (23% of all females) and failure to calve (24% of surviving females). We recommend translocating older females (≥6 years old) because they do not exhibit the reproductive delay and low ORRs of juveniles (<4 years old) or the higher rates of recruitment failure of juveniles and young adults (4–5.9 years old). Where translocation of juveniles is necessary, they should be released into female‐biased populations, where they have higher ORRs. Our study offers the unique advantage of a long‐term analysis across a large number of replicate populations—a science‐by‐management experiment as a proxy for a manipulative experiment, and a rare opportunity, particularly for a large, critically endangered taxon such as the black rhinoceros. Our findings differ from previous recommendations, reinforce the importance of long‐term data sets and comprehensive metrics of translocation success, and suggest attention be shifted from ecological to social constraints on population growth and species recovery, particularly when translocating species with polygynous breeding systems.     

Invasion dynamics of the varnish clam (Nuttallia obscurata): a matrix demographic modeling approach

Marine invaders have become a significant threat to native biodiversity and ecosystem function. In this study, the invasion of the varnish clam (Nuttallia obscurata) in British Columbia, Canada, is investigated using a matrix modeling approach to identify the life history characteristics most crucial for population growth and to investigate population differences. Mark-recapture analyses and field collections from 2003 to 2004 were used to determine individual growth, survival rates, and fecundity for two sites. A multi-state matrix model was used to determine population growth rates and to conduct sensitivity and elasticity analyses. A life table response experiment was also used to determine what life history stage contributed most to observed differences in population growth rates. Population survey data were used in conjunction with the matrix model to determine plausible recruitment levels and to investigate recruitment scenarios. Both populations are currently declining but are likely sustainable because of the pulsed nature of large recruitment events. Survival of larger clams (>40 mm) is the most important for population growth based on elasticity and sensitivity analyses. Adult survival also had the largest influence on observed differences between site-specific population growth rates. The two populations studied differed in recruitment dynamics; one experiencing annual recruitment with higher post-settlement mortality and the other, episodic recruitment and lower post-settlement mortality. The most influential factor for the successful invasion of the varnish clam appears to be survival of the larger size classes. Therefore, any process that decreases adult survival (e.g., predation, commercial harvest) will have the greatest impact on population growth.     

Life history benefits of residency in a partially migrating pond-breeding amphibian

Species with partial migration, where a portion of a population migrates and the other remains residential, provide the opportunity to evaluate conditions for migration and test mechanisms influencing migratory decisions. We conducted a five-year study of two populations of red-spotted newts (Notophthalmus viridescens), composed of individuals that either remain as residents in the breeding pond over the winter or migrate to the terrestrial habitat. We used multistate mark-recapture methods to (1) test for differences in survival probability between migrants and residents, (2) determine if migrants breed every year or skip opportunities for reproduction, and (3) estimate the frequency of individuals switching migratory tactic. We used estimates of life history parameters from the natural populations in combination with previous experimental work to evaluate processes maintaining partial migration at the population level and to assess mechanisms influencing the decision to migrate. Based on capture-recapture information on over 3000 individuals, we found that newts can switch migratory tactics over their lifetime. We conclude that migrants and residents coexist through conditional asymmetries, with residents having higher fitness and inferior individuals adopting the migrant tactic. We found that newts are more likely to switch from residency to migrating than the reverse and males were more likely to remain as residents. Migration differences between the sexes are likely driven by reproduction benefits of residency for males and high energetic costs of breeding resulting in lower breeding frequencies for females. Environmental conditions also influence partial migration within a population; we found support for density-dependent processes in the pond strongly influencing the probability of migrating. Our work illustrates how migration can be influenced by a complex range of individual and environmental factors and enhances our understanding of the conditions necessary for the evolution and maintenance of partial migration within populations.     

Habitat-specific impacts of multiple consumers on plant population dynamics

Multiple consumers often attack seeds, seedlings, and adult plants, but their population-level consequences remain uncertain. We examined how insect and small mammal consumers influenced the demography and abundance of the perennial shrub, bush lupine (Lupinus arboreus). In grassland and dune habitats we established replicate experimental lupine populations in 81-m2 plots that were either protected from, or exposed to, herbivorous voles and granivorous mice (via fencing) and/or root feeding insects (via insecticide treatment). Populations were initiated with transplanted seedlings in 1999 and 2000. We followed the demography of these cohorts, subsequent generations, and the seed bank for 5.5 years. Voles and insects killed many seedlings in dune (1999 only) and grassland (1999 and 2000) habitats. After 2000, insects and voles had minimal effects on seedling or adult survival. Seed predation by granivorous mice, however, greatly depressed seedling recruitment, resulting in lower adult lupine abundance in control plots vs. those protected from rodents. In grasslands, initial effects of voles and insects on seedling survival produced large differences among treatments in adult plant density and the cumulative number of seeds produced throughout the experiment. Differences among grassland populations in seed rain, however, had little influence on the magnitude of seedling recruitment into this habitat. Instead, recruitment out of a preexisting seed bank compensated for the lack of seed production in populations exposed to consumers. Shading by dense adults in plots protected from consumers limited seedling establishment within these populations. Although differences among populations in cumulative seed rain did not influence adult establishment, populations protected from consumers accumulated substantially larger seed banks than controls. These results illustrate how density dependence, habitat-specific seed dynamics, and particular demographic impacts of consumers interact to shape plant population responses to consumers.     

A Comparison of Linear Demographic Models and Fraction of Lifetime Egg Production for Assessing Sustainability in Sharks

Conventional methods for management of data‐rich fisheries maintain sustainable populations by assuring that lifetime reproduction is adequate for individuals to replace themselves and accounting for density‐dependent recruitment. Fishing is not allowed to reduce relative lifetime reproduction, the fraction of current egg production relative to unfished egg production (FLEP), below a sustainable level. Because most shark fisheries are data poor, other representations of persistence status have been used, including linear demographic models, which incorporate life‐history characteristics in age‐structured models with no density dependence. We tested how well measures of sustainability from 3 linear demographic methods (rebound potential, stochastic growth rate, and potential population increase) reflect actual population persistence by comparing values of these measures with FLEP for 26 shark species. We also calculated the value of fishing mortality (F) that would allow all 26 species to maintain an accepted precautionary threshold for sharks of FLEP = 60%, expressing F as a fraction of natural mortality (M). Values of stochastic growth rate and potential population growth did not covary in rank order with FLEP (p = 0.057 and p = 0.077, respectively) and neither was significantly correlated with FLEP. Ordinal ranking of rebound potential positively covaried with FLEP (p = 0.00013), but the relative rankings of some species were substantially out of order. Adopting a sustainable limit of F = 0.16M would maintain all 26 species above the precautionary minimum value of FLEP (60%). We concluded that shark‐fishery and conservation policies should rely on calculation of replacement (i.e., FLEP), and that sharks should be fished at a precautionary level that would protect all stocks (i.e., F< 0.16M). Comparación entre Modelos Demográficos Lineales y la Fracción de Producción de Huevos a lo Largo de la Vida para Estudiar la Sustentabilidad en Tiburones Resumen     

Influence of behavior and mating success on brood-specific contribution to fish recruitment in ponds

One source of uncertainty in predicting the response of populations to exploitation is individual differences within a population in both vulnerability to capture and contribution to population renewal. For species with parental care, individuals engaged in nesting behavior are often targeted for exploitation, but predicting outcomes of this nonrandom vulnerability will depend in part on an understanding of how parental traits are related to potential for brood contribution to the population. Variation in brood-specific contribution to recruitment of largemouth bass (Micropterus salmoides), a fish species with extended parental care, was quantified to determine if differences in mating success, parental care behaviors, and timing of reproduction influenced offspring recruitment. Dependence of these relationships on brood predation was tested in communities that differed in the presence of bluegill, Lepomis macrochirus, an important nest predator. Daily snorkel surveys were conducted in experimental ponds during spring to monitor male spawning and parental care behaviors in populations of largemouth bass. Tissue samples collected from larvae in nests were used to develop brood-specific DNA fingerprints for determining nest origins of fall recruits. Largemouth bass spawning period in bluegill ponds was longer and more variable in duration, with lower, more variable mating success, than in ponds without bluegill. In all populations, only one or two broods provided the majority of recruits, and these were broods produced during the earliest days of spawning by the oldest, largest males. In bluegill ponds, brood contribution from earliest nests also increased with brood size. Earliest nesters were the oldest males, and recruits from these nests were often above average in body size. Offspring needed to be guarded to at least swim-up larval stage to contribute any recruits. Termination of parental protection before offspring were free swimming mainly occurred with broods guarded by smaller males in ponds with brood predators. These age- and size-specific differences in timing of spawning and duration of parental care are consistent with influences of residual reproductive value and energetic constraints on reproductive behavior. Furthermore, these patterns of individual contribution to recruitment imply that fisheries that selectively target either nesting individuals or larger, older males could potentially decrease recruitment at the population scale.     

Sink populations in carnivore management: cougar demography and immigration in a hunted population.

Carnivores are widely hunted for both sport and population control, especially where they conflict with human interests. It is widely believed that sport hunting is effective in reducing carnivore populations and related human-carnivore conflicts, while maintaining viable populations. However, the way in which carnivore populations respond to harvest can vary greatly depending on their social structure, reproductive strategies, and dispersal patterns. For example, hunted cougar (Puma concolor) populations have shown a great degree of resiliency. Although hunting cougars on a broad geographic scale (> 2000 km2) has reduced densities, hunting of smaller areas (i.e., game management units, < 1000 km2), could conceivably fail because of increased immigration from adjacent source areas. We monitored a heavily hunted population from 2001 to 2006 to test for the effects of hunting at a small scale (< 1000 km2) and to gauge whether population control was achieved (lambda < or = 1.0) or if hunting losses were negated by increased immigration allowing the population to remain stable or increase (lambda > or = 1.0). The observed growth rate of 1.00 was significantly higher than our predicted survival/fecundity growth rates (using a Leslie matrix) of 0.89 (deterministic) and 0.84 (stochastic), with the difference representing an 11-16% annual immigration rate. We observed no decline in density of the total population or the adult population, but a significant decrease in the average age of independent males. We found that the male component of the population was increasing (observed male population growth rate, lambda(OM) = 1.09), masking a decrease in the female component (lambda(OF) = 0.91). Our data support the compensatory immigration sink hypothesis; cougar removal in small game management areas (< 1000 km2) increased immigration and recruitment of younger animals from adjacent areas, resulting in little or no reduction in local cougar densities and a shift in population structure toward younger animals. Hunting in high-quality habitats may create an attractive sink, leading to misinterpretation of population trends and masking population declines in the sink and surrounding source areas.     

Demographic Processes Underlying Population Growth and Decline in Salamandra salamandra

Abstract:  Human activity commonly has negative impacts on wildlife. Often, however, only a single element of the life cycle is affected, and it is unclear whether such effects translate into effects on population growth. This is particularly true for research into the causes of global amphibian declines, where experimental research focuses primarily on the aquatic larval stages but theory suggests these stages have only minor importance for population growth. We used data from long-term mark-recapture studies of two natural populations of the salamander Salamandra salamandra to confirm the predictions of population models. One population remained stable (i.e., stationary) throughout the 20 years of the study whereas the other declined to local extinction. We used mark-recapture models to break down population growth rate into its two main components, recruitment and adult survival. Survival of postmetamorphic salamanders was constant over time in the stable population, whereas the declining population was characterized by a decrease in survival and constant recruitment. Population growth was most sensitive to variation in adult survival. Current amphibian research focuses on preadult stages, and researchers assume recruitment is the most important determinant of population growth. This may not be the case. A better understanding of amphibian population dynamics is possible only through the integration of experiments, theory, and data from natural populations. Our results also suggest that amphibian conservation efforts should focus on all stages of the life cycle and their associated habitats.     

Maternal characteristics and the production and recruitment of sons in the eastern kingbird (<Emphasis Type="BoldItalic">Tyrannus tyrannus</Emphasis>)

Sex allocation theory predicts that if variance in reproductive success differs between the sexes, females who are able to produce high-quality young should bias offspring sex ratio towards the sex with the higher potential reproductive success. We tested the hypothesis that high-quality (i.e., heavy) female eastern kingbirds (Tyrannus tyrannus) that bred early in the breeding season would produce male-biased clutches. A significant opportunity for sexual selection also exists in this socially monogamous but cryptically polygamous species, and we predicted that successful extra-pair (EP) sires would be associated with an excess of male offspring. Although population brood sex ratio did not differ from parity, it increased significantly with female body mass and declined with female breeding date, but was independent of the morphology and display (song) behavior (correlates of reproductive success) of social males and EP sires. Male offspring were significantly heavier than female offspring at fledging. Moreover, the probability that male offspring were resighted in subsequent years declined with breeding date, and was greater in replacement clutches, but lower when clutch size was large. Probability of resighting female offspring varied annually, but was independent of all other variables. Given that variance in reproductive success of male kingbirds is much greater than that of females, and that male offspring are more expensive to produce and have a higher probability of recruitment if fledged early in the season, our results support predictions of sex allocation theory: high-quality (heavy) females breeding when conditions were optimal for male recruitment produced an excess of sons.     

Use of fecundity measured directly throughout the breeding season to test a source-sink demographic model

Populations of landbirds (bird species that occupy terrestrial habitats for most of their life cycle) are declining throughout North America (north of Mexico) and Europe, yet little is known about how demography is driving this trend. A recent model of 5 geographically separated populations of Cerulean Warblers (Dendroica cerulea) that was based on within-season sampling of nest survival and fledgling success shows that all populations are sinks (annual reproduction is consistently less than annual adult mortality). I tested this indirect model by directly measuring fecundity (number of female fledglings/female) during the breeding season for 2 years in a Cerulean Warbler population occupying a mature forest in southwestern Michigan (U.S.A.) I determined territories of male birds on the basis of male plumage characters and phases of the nesting cycle (2007) and on uniquely color-banded males (2008). I transferred locations of identified males to topographic maps. I counted all fledglings in territories from May to July each year. The model I tested may apply only to single-brooded species; therefore, I searched the literature to estimate the percentage of single-brooded species in North America. The breeding season of Cerulean Warblers was short- nearly all nests were initiated from mid-May to late June. Nest predation and brood parasitism were primary and rare causes of nest failure, respectively. Significantly fewer Cerulean Warblers fledged from parasitized than from nonparasitized nests. Fledgling survival required to maintain the population size was well above previously published values for Neotropical migrants. Single-brooded species comprise 62% of North American breeding bird species for which the number of broods per year is known; I believe my results may apply to these species. The consistency between identification of populations as sources or sinks on the basis of either model estimates or direct measurements suggests that a demographic model relying on within-season sampling of fecundity is adequate to determine population status of single-brooded avian populations. In addition, on the basis of results of previous studies, annual adult survival rate of the Cerulean Warbler is typical of parulid warblers that are not declining. Thus, low fecundity, here determined with different quantitative methods, can drive status of landbird species with high-observed survival.     

"Cryptic" direct benefits of mate choice: choosy females experience reduced predation risk while in precopula

Despite the central role that female mate choice plays in the production of biological diversity, controversy remains concerning its evolution and maintenance. This is particularly true in systems where females are choosy but do not receive obvious direct benefits such as nuptial gifts that increase a female's survival and fecundity. In the absence of such direct benefits, indirect benefits (i.e., the production of superior offspring) are often invoked to explain the evolution of mate choice. However, females may receive less obvious, or "cryptic," direct benefits, particularly in species with prolonged pre-mating interactions (e.g., precopulatory mate guarding). We assessed the “cryptic” direct benefits of female choice for large male size in two species of freshwater amphipods that do not receive obvious direct benefits. Females paired with large males experienced decreased predation from fish. However, we found that the size of a female’s mate did not affect her predation risk against predatory dragonflies or the harassment she received by single males while paired. Our results demonstrate that even when females receive no traditional direct benefits, female choice for large male size can still provide important direct benefits. Such “cryptic” direct benefits may be common, especially in species with prolonged mating interactions, and are likely important for fully understanding the evolution of mate choice.     

The stress of scramble: sex differences in behavior and physiological stress response in a time-constrained mating system

Iteroparous species maximize lifetime reproductive fitness by balancing current and future reproductive investments. In order to maximize fitness in the face of social or environmental heterogeneity, individuals of the same species may vary in whether they prioritize current reproductive opportunity or sacrifice immediate reproduction in order to prioritize survival and future reproductive potential. Glucocorticoid (GC) secretion plays an important role in mediating this trade-off by promoting behavioral and physiological responses associated with survival, often at the expense of nonessential (e.g., reproductive) functions. We used wood frogs (Lithobates sylvaticus [Rana sylvatica]) to test whether males and females differed in their (a) physiological response (plasma corticosterone [CORT] concentration) to standardized handling stress—a proxy for predation threat—and (b) performance of reproductive behaviors that may enhance their conspicuousness to predators. We also tested whether levels of male competition influenced sex differences in these factors, as more intense competition may require males to devote more time to risky reproductive behaviors. We found that females had lower baseline CORT but exhibited a significantly greater CORT response to a stressor and spent less time performing potentially risky behavior (surface floating) than did males. These sex differences were consistent across different levels of male mating competition. Our results reveal that during breeding, males and females may differentially respond to stressors and perform risk-prone behaviors, despite facing the same extreme breeding constraints, providing new insight into the survival-reproduction trade-off of explosively breeding species.     

Why do most tropical animals reproduce seasonally? Testing hypotheses on an Australian snake

Most species reproduce seasonally, even in the tropics where activity occurs year-round. Squamate reptiles provide ideal model organisms to clarify the ultimate (adaptive) reasons for the restriction of reproduction to specific times of year. Females of almost all temperate-zone reptile species produce their eggs or offspring in the warmest time of the year, thereby synchronizing embryogenesis with high ambient temperatures. However, although tropical reptiles are freed from this thermal constraint, most do not reproduce year-round. Seasonal reproduction in tropical reptiles might be driven by biotic factors (e.g., peak periods of predation on eggs or hatchlings, or food for hatchlings) or abiotic factors (e.g., seasonal availability of suitably moist incubation conditions). Keelback snakes (Tropidonophis mairii, Colubridae) in tropical Australia reproduce from April to November, but with a major peak in May-June. Our field studies falsify hypotheses that invoke biotic factors as explanations for this pattern: the timing of nesting does not minimize predation on eggs, nor maximize food availability or survival rates for hatchlings. Instead, our data implicate abiotic factors: female keelbacks nest most intensely soon after the cessation of monsoonal rains when soils are moist enough to sustain optimal embryogenesis (wetter nests produce larger hatchlings, that are more likely to survive) but are unlikely to become waterlogged (which is lethal to eggs). Thus, abiotic factors may favor seasonal reproduction in tropical as well as temperate-zone animals.     

Climate and predation dominate juvenile and adult recruitment in a turtle with temperature-dependent sex determination

Conditions experienced early in life can influence phenotypes in ecologically important ways, as exemplified by organisms with environmental sex determination. For organisms with temperature-dependent sex determination (TSD), variation in nest temperatures induces phenotypic variation that could impact population growth rates. In environments that vary over space and time, how does this variation influence key demographic parameters (cohort sex ratio and hatchling recruitment) in early life stages of populations exhibiting TSD? We leverage a 17-year data set on a population of painted turtles, Chrysemys picta, to investigate how spatial variation in nest vegetation cover and temporal variation in climate influence early life-history demography. We found that spatial variation in nest cover strongly influenced nest temperature and sex ratio, but was not correlated with clutch size, nest predation, total nest failure, or hatching success. Temporal variation in climate influenced percentage of total nest failure and cohort sex ratio, but not depredation rate, mean clutch size, or mean hatching success. Total hatchling recruitment in a year was influenced primarily by temporal variation in climate-independent factors, number of nests constructed, and depredation rate. Recruitment of female hatchlings was determined by stochastic variation in nest depredation and annual climate and also by the total nest production. Overall population demography depends more strongly on annual variation in climate and predation than it does on the intricacies of nest-specific biology. Finally, we demonstrate that recruitment of female hatchlings translates into recruitment of breeding females into the population, thus linking climate (and other) effects on early life stages to adult demographics.     

Effect of gyrodactylid ectoparasites on host behaviour and social network structure in guppies <Emphasis Type="Italic">Poecilia reticulata</Emphasis>

Understanding how individuals modify their social interactions in response to infectious disease is of central importance for our comprehension of how disease dynamics operate in real-world populations. Whilst a significant amount of theoretical work has modelled disease transmission using network models, we have comparatively little understanding of how infectious disease impacts on the social behaviour of individuals and how these effects scale up to the level of the population. We experimentally manipulated the parasite load of female guppies (Poecilia reticulata) and introduced fish either infected with the ectoparasites Gyrodactylus spp. (experimental) or uninfected (control) into replicated semi-natural populations of eight size-matched female guppies. We quantified the behaviour and social associations of both the introduced fish and the population fish. We found that infected experimental fish spent less time associating with the population fish than the uninfected control fish. Using information on which fish initiated shoal fission (splitting) events, our results demonstrate that the population fish actively avoided infected experimental fish. We also found that the presence of an infected individual resulted in a continued decline in social network clustering up to at least 24 h after the introduction of the infected fish, whereas in the control treatment, the clustering coefficient showed an increase at this time point. These results demonstrate that the presence of a disease has implications for both the social associations of infected individuals and for the social network structure of the population, which we predict will have consequences for infectious disease transmission.     

The evolution of female social relationships in nonhuman primates

Considerable interspecific variation in female social relationships occurs in gregarious primates, particularly with regard to agonism and cooperation between females and to the quality of female relationships with males. This variation exists alongside variation in female philopatry and dispersal. Socioecological theories have tried to explain variation in female-female social relationships from an evolutionary perspective focused on ecological factors, notably predation and food distribution. According to the current “ecological model”, predation risk forces females of most diurnal primate species to live in groups; the strength of the contest component of competition for resources within and between groups then largely determines social relationships between females. Social relationships among gregarious females are here characterized as Dispersal-Egalitarian, Resident-Nepotistic, Resident-Nepotistic-Tolerant, or Resident-Egalitarian. This ecological model has successfully explained differences in the occurrence of formal submission signals, decided dominance relationships, coalitions and female philopatry. Group size and female rank generally affect female reproduction success as the model predicts, and studies of closely related species in different ecological circumstances underscore the importance of the model. Some cases, however, can only be explained when we extend the model to incorporate the effects of infanticide risk and habitat saturation. We review evidence in support of the ecological model and test the power of alternative models that invoke between-group competition, forced female philopatry, demographic female recruitment, male interventions into female aggression, and male harassment. Not one of these models can replace the ecological model, which already encompasses the between-group competition. Currently the best model, which explains several phenomena that the ecological model does not, is a “socioecological model” based on the combined importance of ecological factors, habitat saturation and infanticide avoidance. We note some points of similarity and divergence with other mammalian taxa; these remain to be explored in detail. Received: 30 September 1996 / Accepted after revision: 20 July 1997     

Personality trait differences between mainland and island populations in the common frog (Rana temporaria)

Understanding and predicting species range expansions is an important challenge in modern ecology because of rapidly changing environments. Recent studies have revealed that consistent within-species variation in behavior (i.e., animal personality) can be imperative for dispersal success, a key process in range expansion. Here we investigate how habitat isolation can mediate differentiation of personality traits between recently founded island populations and the main population. We performed laboratory studies of boldness and exploration across life stages (tadpoles and froglets) using four isolated island populations and four mainland populations of the common frog (Rana temporaria). Both tadpoles and froglets from isolated populations were bolder and more exploratory than conspecifics from the mainland. Although the pattern can be influenced by possible differences in predation pressure, we suggest that this behavioral differentiation might be the result of a disperser-dependent founder effect brought on by an isolation-driven environmental filtering of animal personalities. These findings can have important implications for both species persistence in the face of climate change (i.e., range expansions) and ecological invasions as well as for explaining rapid speciation in isolated patches.