Body size variation among mainland populations of the western rattlesnake (Crotalus viridis)

In general, squamate reptiles follow the converse to Bergmann's rule, attaining smaller sizes in cooler environments, whereas other vertebrate groups follow Bergmann's rule, attaining larger sizes in cooler areas. Intensive studies of body size evolution for species of squamates are necessary to understand the processes responsible for this trend. Here I present data on body size variation among mainland populations of the western rattlesnake, Crotalus viridis. This species consists of two well-differentiated phylogenetic clades, therefore all analyses were performed for the C. viridis group as a whole and separately for each of the two clades within the C. viridis group. Although both phylogenetic and nonphylogenetic analyses were performed, the data did not show phylogenetic conservatism, and therefore the nonphylogenetic results are preferred. I found no significant relationships between mean adult female snout-vent length and any of the physical and climatic variables that were examined for the C. viridis group using simple linear regression analysis. Examined separately, I found that individuals of the western clade, C. oreganus, were smaller in cooler and more seasonal environments, whereas individuals of the eastern clade. C. viridis sensu stricto, were larger in cooler and more seasonal areas. Thus, the observed size trends were in opposite directions for the two clades. Multiple regression analysis revealed that seasonality was a stronger predictor of body size variation than was temperature for both clades. The differences in body size trends between these clades may be due to differences in mortality rates among populations.     

Seasonal clutch size variation of multi-brooded bird species: comparisons between breeding season and latitudes

Single- and multi-brooded species of birds differ in their seasonal patterns of clutch size. Single-brooded species start with a maximum clutch size that declines continuously as the season progresses, whereas the clutch sizes of multi-brooded species usually increase to a mid-season maximum peak and then decrease progressively until the end of the breeding season. Previous studies have shown that multi-brooded migrant species present seasonal patterns that are similar to single-brooded species at high latitudes but similar to multi-brooded non-migratory species at lower latitudes. We studied the Greenfinch Carduelis chloris and Goldfinch C. carduelis populations in eastern Spain (Sagunto, Valencia) between 1975 and 2002 to compare seasonal variations in clutch sizes between years with early and late starts to the breeding season. The period over which clutch sizes increase was longer when the breeding season started earlier. The Goldfinch population showed no pattern of initial increase in clutch size when there was a late start to the breeding season: a late start shortens the season giving them less time to breed, and may also coincide with maximum food availability. Thus, the pattern of single-brooded species was observed. In the Greenfinch population, a trend toward the seasonal pattern of single-brooded species was also observed when the following indices were compared: clutch size increase, modal timing, initial slope and timing of maximum clutch size. We have also compared the seasonal patterns of clutch size of both species in eastern Spain with the patterns observed in Britain. Our results show that for both Goldfinches and Greenfinches, the non-migrant southern populations of Sagunto in eastern Spain do not tend towards a more multi-brooded seasonal pattern of clutch size than the migrant Goldfinches of Britain.     

Influence of population density on group sizes in goitered gazelle (Gazella subgutturosa Guld., 1780)

We conducted our study in Ili depression, south-eastern Kazakhstan during 1981–1989 to investigate how group sizes and group class frequencies change with increasing population densities in goitered gazelles. In addition, we compared our study to data on group size and group class frequency of various goitered gazelle populations in Kazakhstan with very variable population densities. We found that mean group size was a more variable index than group class frequency. Population density had some effect on mean group sizes, but the strength of the influence was quite weak, and only in cases where densities of two populations varied more than sevenfold did group sizes start to change. Group class frequency was not correlated with population density at all. The impact of the yearly breeding cycle on group size was bigger than population density. The density-dependent response of goitered gazelle population was curvilinear in fashion, and it may be classified as intermediate between social-dwelling ungulate species, living in large groups and demonstrating continuous (linear) increases of group size with population density and those that are solitary or territorial ungulate species with no relationship between population size and group size, though the goitered gazelle population’s weak response was distinctively closer to the one of solitary ungulate species.     

Seasonal population dynamics in Lucilia eximia (Wiedemann) (Diptera: Calliphoridae)

In this study the seasonal variation of fecundity, wing and tibia sizes were investigated in natural populations of Lucilia eximia (Wiedemann) as an attempt to determine the variations in life history of the species associated to seasonality. Specimens of L. eximia were monthly collected in Botucatu, S?o Paulo, Brazil, during two years and the adult females dissected to estimate fecundity. Body size was estimated by measuring wing and tibia. Fecundity and body size were seasonally analysed. A relatively constant temporal trajectory was found for fecundity, wing and tibia size over twenty-four months. Strong positive correlations between wing and tibia size, fecundity and wing and fecundity and tibia were observed. The maintenance of stable values in L. eximia indicates that it has suffered little seasonal influence over the period analysed. This result confirms the demographics and life history stable profile of the species in spite of seasonal climatic changes observed in the study area.     

Cetaceans of Southeast Alaska: distribution and seasonal occurrence

Aim To assess the distribution, group size, seasonal occurrence and annual trends of cetaceans. Location The study area included all major inland waters of Southeast Alaska. Methods Between 1991 and 2007, cetacean surveys were conducted by observers who kept a constant watch when the vessel was underway and recorded all cetaceans encountered. For each species, we examined distributional patterns, group size, seasonal occurrence and annual trends. Analysis of variance (anova F) was used to test for differences in group sizes between multiple means, and Student’s t‐test was used to detect differences between pairwise means. Cetacean seasonal occurrence and annual trends were investigated using a generalized linear model framework. Results Humpback whales (Megaptera novaeangliae) were seen throughout the region, with numbers lowest in spring and highest in the fall. Fin whale (Balaenoptera physalus) and minke whale (Balaenoptera acutorostrata) distributions were more restricted than that reported for humpback whales, and the low number of sightings precluded evaluating seasonal trends. Three killer whale (Orcinus orca) eco‐types were documented with distributions occurring throughout inland waters. Seasonal patterns were not detected or could not be evaluated for resident and offshore killer whales, respectively; however, the transient eco‐type was more abundant in the summer. Dall’s porpoise (Phocoenoides dalli) were distributed throughout the region, with more sightings in spring and summer than in fall. Harbour porpoise (Phocoena phocoena) distribution was clumped, with concentrations occurring in the Icy Strait/Glacier Bay and Wrangell areas and with no evidence of seasonality. Pacific white‐sided dolphins (Lagenorhynchus obliquidens) were observed only occasionally, with more sightings in the spring. For most species, group size varied on both an annual and seasonal basis. Main conclusions Seven cetacean species occupy the inland waters of Southeast Alaska, with distribution, group size, seasonal occurrence and annual trends varying by species. Future studies that compare spatial and temporal patterns with other features (e.g. oceanography, prey resources) may help in identifying the key factors that support the high density and biodiversity of cetaceans found in this region. An increased understanding of the region’s marine ecology is an essential step towards ensuring the long‐term conservation of cetaceans in Southeast Alaska.     

Ecological correlates to activity and habitat use of two prosimian primates: Eulemur rubriventer and Eulemur fulvus rufus in Madagascar

In this paper, I provide data on the possible effects of group size and seasonal changes in food availability on the activity and habitat use patterns of two species of prosimian primate: the rufous lemur (Eulemur fulvus rufus) and the red-bellied lemur (Eulemur rubriventer). General and subtle seasonal differences were observed between lemur species in (1) group size and composition, (2) activity profiles, and (3) habitat use. Rufous lemur groups were larger (mean = 8 individuals) and contained more adults than red-bellied lemur groups (mean = 3 individuals). The overall degree and distribution of diurnal activity differed between lemur species and varied within species with seasonal changes in food availability. In general, rufous lemurs traveled more often than red-bellied lemurs. During food scarcity, both species increased the amount of time spent feeding, although peak feeding and traveling times differed. Both species also preferred horizontal substrates; however, during food scarcity, red-bellied lemurs used terminal branches more often while feeding than rufous lemurs. In addition, red-bellied lemurs used the lower middle story forest and rested in taller, upper canopy trees more often than rufous lemurs. Differences in activity and habitat use patterns between lemur species were more related to seasonal changes in food availability than overall differences in group size. These behavioral patterns may represent different strategies used by each lemur species to avoid directly competing for similar resources during times of food scarcity. © 1996 Wiley-Liss, Inc.     

Widespread reproductive variation in North American turtles: temperature, egg size and optimality

Theory predicts the existence of an optimal offspring size that balances the trade-off between offspring fitness and offspring number. However, in wild populations of many species, egg size can still vary from year to year for unknown reasons. Here, we hypothesize that among-year variation in population mean egg size of freshwater turtles is partly a consequence of their gonadal sensitivity to seasonal temperatures, a physiological mechanism which principally functions to synchronize reproduction with a favorable time of year. As part of this process, among-year variation in seasonal temperatures modifies the extent of egg follicle development, and this may translate into variation in mean egg size among years (both at the individual and population level). To test this hypothesis, we applied an information-theoretic approach to model relationships between mean egg mass and the temperatures experienced during discrete periods of follicular development in wild populations of three turtle species (Chrysemys picta, Chelydra serpentina, Glyptemys insculpta) over 12 consecutive years. Because follicular development occurs in the fall for C. serpentina and G. insculpta, whereas it occurs both in the fall and spring for C. picta, we expected only fall temperatures would explain egg size variation in C. serpentina and G. insculpta, whereas both fall and spring temperatures would correlate with egg size variation in C. picta. These predictions were upheld. We then compared among-year variation in within-female egg and clutch sizes of each species in order to evaluate whether such variation might still be consistent with some tenets of optimal egg size theory. In all three species, we found that clutch sizes vary more than egg sizes in spite of temperature-induced egg size variation, and this pattern of relatively high clutch-size variation matches theoretical predictions. Future work should explore the roles of direct and indirect (i.e., nutritional) influences of temperature on egg size in natural settings.     

Patterns in the geographic ranges of the world's woodpeckers

We used data on the world's woodpeckers to test for patterns in the geographic distributions of a single group of closely related species. The frequency distribution of woodpecker geographic range sizes is approximately lognormal. Most variation in range sizes is explained by differences between species within genera; that is, range size seems to be an evolutionarily labile trait. The largest woodpecker ranges are found in Eurasia, both when absolute differences are compared and when range size is measured as a proportion of estimated available habitat. Notably, there is a negative relationship between the mean range sizes attained by species in a genus or tribe in South America and the mean ranges attained by species in the same tribe or genus in North America. Large-bodied species tend to be more widely distributed and to live at higher latitudes, but both tendencies disappear if the taxonomic relatedness of species is controlled for. Species living at high latitudes also tend to be more widely distributed. This relationship seems largely due to the effect of North American woodpeckers, which show it even when the taxonomic relatedness of species is controlled. Small continents generally have more woodpecker species than do large ones. Woodpecker geographic range sizes are smaller the more woodpecker species inhabit an area. Species show less overlap in their geographic ranges with species of similar than with species of dissimilar body size. The implications of these results for our understanding of patterns in geographic range sizes are discussed.     

Geographical differences in seasonal clutch size variation in multi-brooded bird species

The seasonal pattern of clutch size variation in birds varies among species. In single-brooded bird species clutch sizes decline continuously with date from an early season maximum. In resident multi-brooded species, clutch sizes first increase to a mid-season maximum and then decrease again. Limited data for multi-brooded migratory birds suggest that clutches in these species also show a continuous decline throughout the season, but it remains to be resolved whether this applies generally and whether migration adds a constraint to the system. We here report on a study of the Eastern Bluebird Sialia sialis conducted using data collected by volunteers across most of its range. In the southern part of its range, where Eastern Bluebirds are largely year-round residents, clutches initially increase, reach a mid-season maximum and gradually decline later in the season. In the northern part of the range, where the majority of Eastern Bluebirds are migratory, clutch sizes decline continuously throughout the season. To determine whether seasonal changes in the clutch size of multi-brooded species are determined by migratory behaviour of the population we compared our results with published data on the Dunnock Prunella modularis and conclude that in multi-brooded species migratory behaviour is not a sufficient condition for a continuous seasonal decline in clutch size, but it could be a necessary one. We propose two mechanisms for the pattern observed among multi-brooded migrants: (i) a time or energy cost of migration, and (ii) a more abrupt increase in seasonal resources in spring at more northerly latitudes.     

Seasonal changes in spermatogenesis in the blue fox (Alopex lagopus), quantified by DNA flow cytometry and measurement of soluble Mn2+ -dependent adenylate cyclase activity

The testes of the blue fox (Alopex lagopus) showed marked seasonal variations in size. Testicular weight and volume increased rapidly during January and February to reach maximal values by the beginning of the breeding season (approximately 15 March). During May and June the weights and volumes of the testes declined gradually to the quiescent state which lasted from July until October. Quantitation by DNA flow cytometry of the seasonal changes in the relative numbers of haploid (1C), diploid (2C) and tetraploid (4C) cell numbers in the testis showed that the increase in testis size from December to February was associated with a rapid expansion of the haploid cell compartment as spermatogenesis resumed. In addition, an increase in number of more mature cell types within the haploid cell population was observed over a 2-month period before the breeding season. The decline in testicular size from the middle of April until October was associated with a reduction in both the absolute and relative sizes of the haploid and tetraploid cell populations and a concomitant increase in the relative numbers of diploid cells. Measurements of the activity of the soluble Mn2+ -dependent adenylate cyclase revealed seasonal variations that closely paralleled those of the haploid cell population, indicating that, as in other species, the enzyme may be associated with maturing germ cells.     

Geographical range size of tropical plants influences their response to anthropogenic activities

Biodiversity patterns may be influenced by the species geographical range sizes, but this is rarely shown. We used a highly replicated and large-scale study in coastal Ecuador to determine for the first time the importance of latitudinal range size of plant species in their response to land-use activities. We examined herbaceous plant communities of five land-use types with decreasing anthropogenic disturbance (from the most intervened rice and pasture to the less intervened managed agroforest, abandoned agroforest, and forest) in a low and a high impact human-dominated landscape. All species were classified in four latitudinal range size quartiles, from the 25% species with the narrowest to the 25% with the widest range size. We found notable differences between patterns of total species richness and those of individual range size quartiles. Whereas total species richness was higher in more intervened land-use types, percentages of narrow-ranged species were significantly higher in less intervened land-use types. In contrast, percentages of wide-ranging species were higher in more intervened land-use types. Hence, responses of plant species to human activities were influenced by traits that determine their range sizes. An analysis of floristic similarity between land-use types revealed that narrow-ranged species were mainly preserved in forest fragments, but the other land-use types supported many unique narrow-ranged species and therefore made an important contribution to their preservation at the landscape level. Conservation efforts should combine protection of natural habitats with strategies to maintain a diversity of low-intensity land-use types, looking for win-win solutions or trade-offs between biodiversity conservation and human welfare in human-dominated landscapes.     

Does size matter? Relating consumed prey sizes and diet composition of otters in South Iberian coastal streams

We analysed seasonal changes in the sizes of prey [grey mullets (Mugilidae), flatfish (Soleidae), eelAnguilla anguilla and crayfishProcambarus clarkii] consumed by ottersLutra lutra Linnaeus, 1758 in a Mediterranean-climate coastal area and relate them to the frequencies of occurrence of each prey species in otter diet. The sizes of over 1500 otter prey were estimated from measurements of key pieces found in 814 otter spraints, which were collected in lower stream stretches in a sandy coastal area. Clear relationships between mean prey size and frequency of occurrence were observed for the four prey types. These relationships were positive for grey mullets, flatfish and crayfish (ie they occurred more in otter diet when more large individuals were predated), but was negative in the case of eels. Results suggest that these patterns could be related to seasonal changes in habitat use. Previous works in the study area showed that otter concentrate its predation efforts in freshwater stream stretches during spring and summer, when more and larger crayfish are available. Freshwater stretches have neither grey mullets nor flatfish, while eels are larger and scarcer there than near streams’ mouths, where crayfish is absent.     

Turbellarian assemblages in freshwater lagoons in southern Brazil

Turbellarians, which typically feed on bacteria, algae, rotifers, oligochaetes, dipteran larvae, microcrustaceans, and other organisms, are abundant in diverse types of wetlands. Despite their importance, abundance, and species richness in freshwater environments, turbellarians are seldom considered in studies on biodiversity. We analyzed the structure of turbellarian communities in shore areas of three categories of permanent wetlands classified according to their perimeter as small, intermediate, and large during an annual cycle. In total, 1847 turbellarians were collected representing 42 species and 15 genera, from the orders Catenulida, Macrostomida, Lecithoepitheliata, Rhabdocoela, and Tricladida. Sixteen species were common to the three categories of wetlands, whereas nineteen species were unique to a particular category. Species composition varied among wetlands of different sizes; small, intermediate, and large wetlands had different dominant species. We found seasonal differences in community composition over the year, but no significant differences in mean values of observed species richness among wetlands with different sizes and among seasonal samples. The estimated species richness was, however, higher in the small wetlands, followed by the large and intermediate wetlands. In the summer, abundance was significantly lower in the small water bodies than in the intermediate and large bodies of water. Our results reinforce the need for conservation of wetlands of different sizes.     

Climate‐driven elevational variation in range sizes of vascular plants in the central Himalayas: A supporting case for Rapoport's rule

A fundamental yet controversial topic in biogeography is how and why species range sizes vary along spatial gradients. To advance our understanding of these questions and to provide insights into biological conservation, we assessed elevational variations in the range sizes of vascular plants with different life forms and biogeographical affinities and explored the main drivers underlying these variations in the longest valley in China''s Himalayas, the Gyirong Valley. Elevational range sizes of vascular plants were documented in 96 sampling plots along an elevational gradient ranging from 1,800 to 5,400 m above sea level. We assessed the elevational variations in range size by averaging the range sizes of all recorded species within each sampling plot. We then related the range size to climate, disturbance, and the mid‐domain effect and explored the relative importance of these factors in explaining the range size variations using the Random Forest model. A total of 545 vascular plants were recorded in the sampling plots along the elevational gradient. Of these, 158, 387, 337, and 112 were woody, herbaceous, temperate, and tropical species, respectively. The range size of each group of vascular plants exhibited uniform increasing trends along the elevational gradient, which was consistent with the prediction of Rapoport''s rule. Climate was the main driver of the increasing trends of vascular plant range sizes in the Gyirong Valley. The climate variability hypothesis and mean climate condition hypothesis could both explain the elevation–range size relationships. Our results reinforce the previous notion that Rapoport''s rule applies to regions where the influence of climate is the most pronounced, and call for close attention to the impact of climate change to prevent species range contraction and even extinction due to global warming.     

Ecological comparisons of robber fly species (Diptera: Asilidae) coexisting in a neotropical forest

Summary Data regarding seasonal abundance, microhabitat preference, and diet were collected over 3 field seasons for adults of 15 robber fly species on Barro Colorado Island, Panama. The species comprised 2 distinct thermal guilds; light-seeking (hereafter LS) species foraged in sunlit areas, while shade-seeking (SS) species foraged only in deep shade. All species were rare during the dry season. During the rainy months, most SS species had flight periods of 2–3 months, and no temporal segregation was apparent. In contrast, most LS species had flight periods of only 4–6 weeks, and a distinct sequence of occurrence was evident during 2 years of censusing. Most SS and LS species displayed a pronounced specificity for perches of a particular substrate type. However, the ranges of perching heights utilized varied considerably among species. Dietary comparisons revealed that mean and maximum prey sizes increased with increasing robber fly size, while minimum prey sizes were constant. Robber fly species <20 mg fed primarily upon nematocerous Diptera, whereas larger species generally fed upon a wide variety of prey types. For each thermal guild, the actual mean overlap for a particular niche dimension was compared to mean overlaps generated by randomly assigning species to thermal guilds. No significant differences from the random null hypothesis were found for the SS guild. However, niche complementarity between dietary and spatial overlaps and dietary overlap was apparent among the 5 large LS species.     

王朗国家级自然保护区岩羊集群结构及季节变化

岩羊(Pseudois nayaur)是青藏高原常见的食草动物,但对边缘分布区的种群了解较少.2018年6月到2019年8月,我们利用红外相机在四川王朗国家级自然保护区对岩羊的集群结构、特征及其季节变化进行了描述与分析.结果显示:调查共记录到岩羊1 921群次,共计6 623只次.按照性别和年龄组,可将岩羊集群划分为混...     

Study of the chorion of seasonal and non-seasonal Africa and Neotropical oviparous Cyprinodontiforme fishes

The structure of the chorion from the fertilized eggs of eight species of killifish, four Nothobranchiidae from tropical Africa and four Rivulidae species from South America and an Asian Cyprinidae (zebrafish), were investigated for possible structural similarities by transmission and scanning electron microscopy. The study of the possible chorion variations between seasonal fishes inhabiting temporary pools of tropical regions and other, non-seasonal species revealed variations in the several types of complex adornments found on the external chorion surface when compared with the less complex chorion of the zebrafish. The inner structure of the chorion of these killifish species comprises alternating electron dense and clear strata, with the number of strata varying by species as well as the thickness of the chorion. No obvious phylogenetic or ecological relationships were observed either between the chorion adornments and the inner organization of the chorion or between the seasonal and non-seasonal species. It is advised to perform studies with a larger sample of species that could show a relation either from ecological or phylogenetic parameters.     

Nest groups of wild bonobos at Wamba: selection of vegetation and tree species and relationships between nest group size and party size

We examined the location of nest groups, spatial distribution of nests within a nest group, and attributes of individual nests of wild bonobos at Wamba, Democratic Republic of Congo. We also examined the seasonal factors influencing nesting behavior and compared the nest group size with the 1 hr party size during daytime. We defined a nest group to be a cluster of nests that were built in the same evening and found within 30 m from the other nearest nest. Examination of the largest gap within a nest group suggested that 30 m was an acceptable cutoff value. Monthly rainfall or fruit abundance did not significantly influence the monthly mean nest group size. Nests were built in the swamp forest for as many as 13% observation days, suggesting the need for reevaluation of the use of swamp forest by bonobos. The use of swamp forest was influenced not by seasonal rainfall or fruit abundance, but by the fruiting of specific species. Preferred tree species for building nests accounted for 19.8% of standing trees, which suggested that the selection of sleeping sites was not largely restricted by the distribution of specific species. The mean 1 hr party size was almost identical through the day and was similar to the mean nest group size. Parties of bonobos sometimes split into smaller nest groups, especially when feeding on non‐preferred fruits during fruit scarcity. By contrast, when feeding on preferred fruits while ranging in large parties, they often aggregated to form even larger nest groups. When sleeping in small‐ or middle‐sized nest groups, they tended to aggregate the next morning. These tendencies may reflect the gregarious nature of bonobos who prefer to range or sleep together as far as circumstances allow. Am. J. Primatol. 72:575–586, 2010. © 2010 Wiley‐Liss, Inc.     

Changes in nonspecific immunity factor in some Equidae, Camelidae and Capridae species

Plasma lysozyme activity changed during the 2 years of the investigation period. In some species (e.g. ponies and goats) there are only seasonal fluctuations with the highest levels during summer and autumn in ponies and during winter in goats. In two Camelidae species, guanacos and llamas, only, downward trends were found, whereas in donkeys, both types of changes: seasonal variations with the maximal level during autumn and an upward long-term trend were found.     

A note on group sizes of oribi (Ourebia ourebi, Zimmermann, 1783) from two contrasting sites in Zambia, with and without predation

The oribi (Ourebia ourebi, Zimmermann, 1783) is a small species of antelope widely represented across open grasslands of sub‐Saharan Africa. Although largely territorial, differences in group size and mating systems (monogamy/polygamy), have been linked to habitat conditions and population density. At high population densities, additional males are recruited to assist in territorial defence. Other factors which might impact on group size, include predation threat, especially at low population densities. In this study (1995–98), we recorded group sizes of oribi across two contrasting areas (c. 300 km² each) of Zambia with (Kafue), and without (Bangweulu), large vertebrate predators. A total of 412 groups was recorded across the two sites, with mean sizes of 2.31 (n = 217) at Bangweulu and 2.33 (n = 195) at Kafue. The modal group size was two throughout (range 1–6). Groups were more variable in size and sex composition at Kafue than at Bangweulu, but there was no significant difference in estimates of population density between sites. Mean estimates were 2.02 and 1.90 km^?2 for Bangweulu and Kafue, respectively. Sex ratios (males : females) were biased towards females (1 : 1.72 and 1 : 3.55, respectively) at both sites. Adult males were particularly scarce at Kafue. However, values of density and group size, were both unexceptional for the species. Whilst a modal group size of two is consistent with a monogamous mating system, variations in group composition and size were more difficult to explain. Adult females appeared to retain or recruit additional animals to groups, irrespective of the needs of territorial defence. This suggests that vigilance is an important function of groups on large open plains, especially in the presence of predators. By extension, reversed sexual dimorphism in oribi could reflect increased selection for vigilance duties in females. Predators had no apparent effect on population density, but oribi groups were more variable in size and sex composition in their presence. Adult males may be particularly vulnerable to predation when defending territorial borders at low population densities. However, further work is needed to define group dynamics in this species and to identify causes of mortality amongst adult males.