Observations on the food of Antedon bifida (Echinodermata: Crinoidea)

The gut contents of specimens of the temperate crinoid Antedon bifida (Pennant) were collected from Dalkey, County Dublin, at monthly intervals for 1 yr, and occasionally from other areas around the Irish coast. Quantitative analysis showed the main components to be detritus (65%), plankton (18%) and inorganic particles (17%). This composition was remarkably constant throughout the year and at all locations, the main variation being in the plankton which was at a minimum in February (4%) and reached a maximum in August (36%). Food particle collection appeared to be nonselective within the appropriate size group, the essential feature of a particle being that it would fit into the ambulacral groove. Fragile planktonic diatoms and dinoflagellates, which occurred in blooms in summer and autumn, were digested by A. bifida, but more robust diatoms passed intact through the gut. Histochemical investigations indicate that the bulk of the detritus is refractory, and it is suggested that its associated micro-flora and micro-fauna may be a major source of nutriment for this crinoid.     

Diet of four species of deep-sea isopods (Crustacea: Malacostraca: Peracarida) in the South Atlantic and the Southern Ocean

The food of four species of asellote isopods (Crustacea, Malacostraca), Haploniscus rostratus, Haploniscus unicornis, Acanthocope galatheae and Betamorpha fusiformis, was evaluated by analysis of their gut contents. The isopods were sampled at several stations on the abyssal plains of Guinea Basin, Angola Basin and Cape Basin (southeast Atlantic), the Weddell Sea abyssal plain and the Antarctic continental slope during the DIVA and ANDEEP expeditions in 2000, 2001 and 2005. While all species had mineral particles in their guts and mucus material was the most frequent food item, the remaining gut contents differed among species. Betamorpha fusiformis fed mostly on phytodetritus, especially in the Southern Ocean basins and ingested along with it whole calcareous foraminifers. Acanthocope galatheae showed some differences in gut contents between basins, but in the Guinea Basin, the contents were to a large extent stercomata, i.e., waste pellets of soft-walled foraminifers, i.e., the Komokiaceae. Indications were that the haploniscids were feeding on detritus and agglutinating foraminifers (stercomata). This indicates spatial differences in food availability for this diverse group of deep-sea isopods and the importance of poorly known foraminiferal groups, like the Komokiaceae, as a food source in the deep sea.     

Lipid compositions,water contents,swimbladder morphologies and buoyancies of nineteen species of midwater fishes (18 myctophids and 1 neoscopelid)

Lipid compositions, water contents, swimbladder morphologies and specific gravities were studied for 19 species of oceanic midwater fishes, chiefly myctophids (lanternfishes) collected offshore from Oregon and California from 1975–1979. Three groups of species were recognizable. The first group had bodies low in both lipid and water content; they were denser than seawater, regardless of swimbladder morphology, which varied from absent through non-inflated to inflated. The second group had bodies low in lipid but high in water content; they were neutrally buoyant, evidently because of their high content of water, although their swimbladders were never inflated. The majority of the members of the aforementioned groups contained higher proportions of triglycerides than wax esters. The third group had bodies with high lipid but low water content; those with high triglyceride content had swimbladders ranging from non-inflated to (less commonly) inflated, were denser than seawater, and the lipid percentage of their body weights varied both with size and season, indicating that triglycerides function mainly as an energy store. The adults of species in the third group with a higher content of wax esters than triglycerides lacked inflated swimbladders, were neutrally buoyant, and the lipid percentage of their body weights was relatively constant with size and season, indicating that wax esters permit these fishes to attain neutral buoyancy in seawater.     

A non-motorized dye ejector for visualization of flow in situ and its use with coral reef crinoids

We describe a portable, non-motorized device for delivering a tracer dye into seawater under field conditions. Dye is ejected at a constant flow rate over a period of tens of minutes. The ejector works in a wide range of ambient pressures without external energy requirements. The flow rate is adjusted simply by varying the length of the delivery tube. The dye streams permitted observations of the upcurrent and downcurrent flow regimes for a filter-feeding crinoid (Comanthus bennetti) living at a depth of 8 m on a coral reef. The results indicate that the crinoid may enhance the rate of particle capture by changing the scale of turbulence in the water passing through the mesh of the filtration fan.     

Characterizing the resident, fermentative microbial consortium in the hindgut of the temperate-zone herbivorous fish, Hermosilla azurea (Teleostei: Kyphosidae)

The zebraperch, Hermosilla azurea Jenkins and Evermann, a warm-temperate marine fish species with a strictly macroalgal diet, has a relatively long digestive tract with an enlarged hindgut and an associated blind caecum (HC). In zebraperch sampled off Santa Catalina Island, California (33°19′42′′N; 118°18′37′′W) in years 1995 through 2001, direct cell counts, gut epithelium assessment of bacterial attachment, and short-chain fatty acid (SCFA) analyses verified that the zebraperch HC possesses a dense and morphologically diverse, fermentative microbiota. Bacterial cell counts and morphological diversity were significantly higher in HC contents compared to anterior gut regions, suggesting that microbial populations were growing along the digestive tract. Similarly, electron micrographs of the HC epithelium revealed attached microbes, further supporting the possibility that these organisms constitute resident microbiota. Five different SCFAs were detected in all three regions of the digestive tract, but levels were up to three times greater in HC contents. Acetate was consistently the prevailing SCFA in all gut regions. Sequence analysis of bacterial 16S rDNA was used to identify predominant bacterial groups in HC contents. Of the seven main bacterial types identified, Enterovibrio spp. were the dominant bacteria in HC contents followed by species of Bacteroides,Faecalibacterium, and Desulfovibrio. Taken together, our findings show that the zebraperch HC harbors a consortium of microbes that appears to assist in the breakdown of algal polysaccharides in the herbivorous diet of the fish.     

Sex-based differences in density-dependent sociality: an experiment with a gregarious ungulate

For animals living in natural or semi-natural settings, empirical data on how sociality changes in response to increasing population density are few, especially with respect to true conspecific density and not group size. However, insight into this line of research may be far-reaching--from understanding density dependence in sexual selection to improving models of disease transmission. Using elk (Cervus elaphus Linnaeus) held in enclosures, we conducted sex-stratified experiments to test how the frequency of dyadic pairings (interaction rate) and their quality (duration) responded to manipulations in exposure to density. Using proximity-logging radio collars we recorded when and for how long individuals shared a space within 1.4 m of each other. As predicted, males increased their interaction rate as density increased. Female interaction rates, however, increased initially as density increased but soon declined to become indistinguishable from rates at low density. Females interacted for longer periods at medium densities, whereas male interaction length clearly decreased as density increased. We highlight a sexually dichotomous, density-dependent response in sociality that has yet to be reported. In addition to furthering our understanding of sociobiology (e.g., implications of time constraints presented by density on dyadic interactions), our results have implications for managing communicable disease in gregarious species of livestock and wildlife.     

Plant Reproductive Traits as a Function of Local, Regional, and Global Abundance

Most ecological generalizations stem from the study of common organisms, but most species are rare. There are a number of reasons to expect that rare and common species may differ consistently in their characteristics, with possible implications for conservation. Past analyses of this issue, however, have generally considered only a single measure of rarity and have not corrected analyses for the lack of independence in the traits of related species. We compared several reproductive characteristics of Mediterranean annual crucifers as a function of their global range, regional abundance, and local population density in Israel, making use of independent contrast methods to correct for the phylogenetic relationships of the species involved. We found plants growing at low local density to be disproportionately likely to be self-compatible. Petal length and floral depth (sepal length) were correlated with breeding system but showed significant interaction effects between breeding system and abundance. Floral longevity was inversely related to abundance and also showed significant interaction effects between breeding system and abundance. Overall, rare species tended to display more extreme values for floral traits than did common plants with the same breeding systems; they had unusually large and deep flowers if self-incompatible and unusually small, shallow ones if self-compatible.     

Copulation duration and sperm competition in white-faced dragonflies (Leucorrhinia intacta; Odonata: Libellulidae)

Summary In many odonates, females mate with more than one male while laying a single clutch of eggs. We studied paternity of eggs laid by remated females of Leucorrhinia intacta, a small libellulid dragonfly, at a pond near Syracuse, NY, USA. The probability of a female remating is a function of male density on the pond. The length of copulations differs considerably among males active on the study pond at the same time. Much of this variation was correlated with differences in mating tactics of the males; copulations by males that stayed on their territories during copulation were shorter than those by other males (Fig. 2). Eggs collected from females mated to irradiated, sterile males and to free-living, fertile males indicated that the average paternity expectation was higher for long than for short copulations, and that the variance in paternity expectation was lower for long than for short copulations. Some possible causes of the high variation in paternity at low copulation durations and possible reasons for differences in copulation duration between male mating tactics are discussed.     

Reproductive biology of the common antarctic crinoid Promachocrinus kerguelensis (Echinodermata: Crinoidea)

Histological examinations of the gonads of Promachocrinus kerguelensis Carpenter, 1888 collected in November 1984 from McMurdo Sound, Antarctica, revealed ripe ovaries containing three distinct size classes of oocytes: 20 to 90, 91 to 150 and 151 to 180 m diameter as well as ova. Testes were ripe, and filled with mature sperms. There was marked inter-pinnule and inter-individual synchrony of gametogenesis and spawning is probably synchronous, at least within the sub-population sampled. The number of eggs per pinnule is allometrically related to body size. Body weight and arm length are positively correlated with total egg number. Annual fecundity was estimated to be 29 000 eggs (full-grown oocytes and ova) for a crinoid weighing 53.6 g wet weight with an arm length of 16.3 cm. The eggs float. Female genital pinnules contain more lipid and energy than male pinnules. Reproductive output and effort (23.4 to 65.2 kJ and 26 to 38%, respectively) increase with body size and are similar to values reported for other echinoderms with lecithotrophic larvae. The greatest overall energetic contribution is represented by the arms and genital pinnules. Spawning in P. kerguelensis probably occurs in November and December within McMurdo Sound. If development is slow, as in all other antarctic echinoderms studied to date, then settlement most likely occurs 2 to 3 mo later, after the short summer period of high phytoplankton productivity. High fecundity, a pelagic mode of reproduction, and the high probability of a slow rate of development may contribute to the wide geographic distribution of this common antarctic crinoid.     

Wood-based diet and gut microflora of a galatheid crab associated with Pacific deep-sea wood falls

Wood falls in the deep sea have recently become the focus of studies showing their importance as nutrients on the deep-sea floor. In such environments, Crustaceans constitute numerically the second-largest group after Mollusks. Many questions have arisen regarding their trophic role therein. A careful examination of the feeding appendages, gut contents, and gut lining of Munidopsis andamanica caught with wood falls revealed this species as a truly original detritivorous species using wood and the biofilm covering it as two main food sources. Comparing individuals from other geographic areas from substrates not reported highlights the galatheid crab as specialist of refractory substrates, especially vegetal remains. M. andamanica also exhibits a resident gut microflora consisting of bacteria and fungi possibly involved in the digestion of wood fragments. The results suggest that Crustaceans could be full-fledged actors in the food chains of sunken-wood ecosystems and that feeding habits of some squat lobsters could be different than scavenging.     

Plasticity in foraging strategies of inshore birds: how Little Penguins maintain body reserves while feeding offspring

Breeding animals face important time and energy constraints when caring for themselves and their offspring. For long-lived species, life-history theory predicts that parents should favor survival over current reproductive attempts, thus investing more into their own maintenance than the provisioning of their young. In seabirds, provisioning strategies may additionally be influenced by the distance between breeding sites and foraging areas, and offshore and inshore species should thus exhibit different strategies. Here, we examine the provisioning strategies of an inshore seabird using a long-term data set on more than 200 Little Penguins, Eudyptula minor. They alternated between two consecutive long and several short foraging trips all along chick rearing, a strategy almost never observed for inshore animals. Short trips allowed for regular provisioning of the chicks (high feeding frequency and larger meals), whereas long trips were performed when parent body mass was low and enabled them to rebuild their reserves, suggesting that adult body condition may be a key factor in initiating long trips. Inshore seabirds do use dual strategies of alternating short and long trips, but from our data, on a simpler and less flexible way than for offshore birds.     

Biochemical and energetic composition of bathyal echinoids and an asteroid,holothuroid and crinoid from the Bahamas

The biochemical and energetic composition of body components of ten species of bathyal echinoids, and an asteroid, a holothuroid and a stalked crinoid were determined from individuals sampled from a variety of deep-water sites near the Bahamas (north Caribbean Sea) in October 1988. When compared with other studies of echinoderms, no geographic- or depth-related differences in biochemical or energetic composition were found. Body-wall tissues were composed primarily of skeletal material (mineral ash), but were comparatively high in organic material in the echinothuriid echinoids, and the asteroid and holothuroid. Gut tissues and pyloric cecae had high levels of lipid and protein, indicating their potential role in nutrient storage. Body-wall tissues were generally low in energy, but were highest in the echinoidsAraeosoma belli (7.7 kJ g^–1 dry wt) andSperosoma antillense (8.0 kJ g^–1 dry wt), the asteroidOphidiaster alexandri (8.9 kJ g^–1 dry wt), and the holothuroidEostichopus regalis (13.1 kJ g^–1 dry wt). Energy levels of gut and pyloric cecal tissues were two to three times higher than those of body-wall tissues. Total somatic tissue energy values varied greatly among species, ranging from 1.5 kJ in the echinoidAspidodiadema jacobyi to 142.1 kJ inE. regalis. As the bathyal echinoderms examined in this study occur in great abundance, they represent a significant reservoir of organic and inorganic materials and energy in deep-water benthic systems.     

Diet of anguilloid larvae: leptocephali feed selectively on larvacean houses and fecal pellets

Gut contents of 234 leptocephali comprising eight species of eels were examined from five families (Congridae, Muraenidae, Muraenesocidae, Nettastomatidae, and Ophichthidae). The larvae belonged to developing leptocephalus (215 specimens) and the early metamorphic stage (19 specimens). Visible gut contents were recognized in 111 individuals among the eight species, regardless of developmental stage. Two kinds of visible objects, transparent (0.4 to 1.2 mm) and opaque (20 to 380 m), were found in the gut of leptocephali. From their morphological characteristics, the former and the latter were identified as larvacean houses and zooplankton fecal pellets, respectively. Furthermore, most fecal pellets in the gut were identified as oikopleurid larvacean's fecal pellets. No trace of the many other phytoplankton or zooplankton, which were found with leptocephali in the ambient waters and could be suitable-size food, was found in the gut of any leptocephalus. On the basis of the importance of larvecean houses in the diet of several species of leptocephalus larvae, it is proposed that the peculiar, large, fang-like teeth of leptocephali are used for feeding, and evolved to pierce and grasp the mucous houses of larvaceans.     

Trophic strategies of euphausiids in a low-latitude ecosystem

Vertical distribution, diet, and morphology of adults were examined in 27 species of euphausiids occurring in the upper 1000 m in the eastern Gulf of Mexico. Vertical distribution patterns were similar to those found in the central ocean gyres and oceanic equatorial waters of the Atlantic, Indian and Pacific Oceans. Most species migrated vertically from their daytime depths of 300 to 600 m to the upper 300 m at night. Exceptions were the non-migrating species of Stylocheiron, which remained in the epipelagic zone day and night, and Nematobrachion boopis, which remained in the mesopelagic zone. Based on gut-contents analysis, the Gulf euphausiids were largely zooplanktivorous, with cyclopoid and calanoid copepods being the most common items in stomachs. ostracods were especially common in the stomachs of Thysanopoda spp. and phytoplankton in the guts of Euphausia spp. Nearly every species' diet contained a considerable amount of olive-colored debris, which may have been marine snow generated in the epipelagic zone. Cluster analysis grouped the euphausiids into nine diet guilds. Euphausiids with a generalized morphology (i.e., spherical eyes, uniform thoracic appendages) tended to group together and demonstrated little variety in stomach contents among species. Euphausiids with a specialized morphology (i.e., bilobed eyes, elongate thoracic appendages) showed considerable variety in stomach contents among species, and several species had diets that were highly specific. Many of the species that had similar gut contents fed on prey of different sizes, as indicated by the width of the calanoid copepod mandibles found in stomachs. Principal-components analysis of seven morphological characters yielded species groups that were similar, but not identical, to those generated by cluster analysis of stomach contents data. We inferred from this that morphological characters partly determine diet, but that behavior is also important. Using the 20 most abundant species and 3 niche parameters, we attempted to identify the degree of separation among euphausiids based on the level of overlap in vertical distribution and diet composition, and on differences in mean prey size. Overlap of <60% in vertical distribution or diet composition was considered to indicate distinction of that parameter. Of 190 total species pairs, only 4 pairs did not demonstrate niche separation in at least one of these categories. We found that differences in these niche parameters were greatest among species with a specialized morphology and least among species that were morphologically generalized.     

Fatty acid trophic markers elucidate resource partitioning within the demersal fish community of South Georgia and Shag Rocks (Southern Ocean)

Fatty acid analysis was used to study the trophic ecology of 10 demersal fish species in the South Georgia region. Principal component analysis grouped the species into three general clusters, revealing resource partitioning between species. Two groups were characterised by large proportions of either monounsaturated or polyunsaturated fatty acids, separating species according to their predominant feeding habitat. The third group showed fatty acid signatures overlapping with either or both of the previous two groups, suggesting a more opportunistic feeding behaviour for these species. Intraspecific comparisons furthermore revealed dietary variability with size, year and geographical location in several species. Mackerel icefish (Champsocephalus gunnari) in particular showed inter-annual differences in muscle lipid concentrations closely linked to prey availability with low lipid contents found in years of low krill (Euphausia superba) abundance. Despite the intraspecific differences the majority of species could be easily distinguished from each other, which indicates the utility of this method in the dietary analysis of higher predators.     

Day length,latitude and behavioural (in)flexibility in baboons<Emphasis Type="Italic"> (Papio cynocephalus ursinus</Emphasis>)

Annual cycles in day length are an important consideration in any analysis of seasonal behaviour patterns, since they determine the period within which obligate diurnal or nocturnal animals must conduct all of their essential activities. As a consequence, seasonal variation in day length may represent an ecological constraint on behaviour, since short winter days restrict the length of the time available for foraging in diurnal species (with long summer days, and thus short nights, a potential constraint for nocturnal species). This paper examines monthly variation in activity patterns over a 4-year study of chacma baboons (Papio cynocephalus ursinus) at De Hoop Nature Reserve, South Africa. Time spent feeding, moving, grooming and resting are all significant positive functions of day length, even before chance events such as disease epidemics and climatically mediated home range shifts have been accounted for. These results provide strong support for the idea that day length acts as an ecological constraint by limiting the number of daylight hours and thus restricting the active period at certain times of year. Day length variation also has important implications across populations. Interpopulation variation in resting time, and non-foraging activity in general, is a positive function of latitude, with long summer days at temperate latitudes apparently producing an excess of time that cannot profitably be devoted to additional foraging or social activity. However, it is the short winter days that are probably of greatest importance, since diurnal animals must still fulfil their foraging requirements despite the restricted number of daylight hours and elevated thermoregulatory requirements at this time of year. Ultimately this serves to restrict the maximum ecologically tolerable group sizes of baboon populations with increasing distance from the equator. Seasonal variation in day length is thus an important ecological constraint on animal behaviour that has important implications both within and between populations, and future studies at non-equatorial latitudes must clearly be mindful of its importance.     

Dealing with urban noise: vermilion flycatchers sing longer songs in noisier territories

In noisy conditions, several avian species modulate their songs in amplitude and in the temporal or frequency domains, presumably to improve communication. Most studies on how passerine birds perform such adjustments have been carried out in oscines, a group well known for the importance of learning in the development of their songs. On the other hand, suboscines, in which learning appears to have little influence on the development of their songs, have been largely neglected. We evaluated song adjustment to noise in the vermilion flycatcher (Pyrocephalus rubinus), a suboscine bird. We conducted song recordings and noise measurements at several territories within Mexico City during the length of the dawn chorus. Males living in noisier places sang long songs, while those males inhabiting quieter places sang both short and long songs. We also found evidence of individual song plasticity, as males sang less versatile songs (i.e., songs with more introductory elements) later in the morning when noise levels were higher. This individual shift in song seems to be more associated to time of the day rather than to the observed rise in noise. However, we cannot discard an effect of noise, which should be evaluated with an experiment. We discuss our results in the context of other studies with oscine passerines and other taxa and consider implications for signaling in intra- and intersexual contexts.     

Patterns of growth,energy utilization and reproduction in some meso- and bathypelagic fishes off Southern California

We have studied growth, energy use and reproduction in 4 mesopelagic fishes and 5 bathypelagic fishes living off Southern California (USA). All of the mesopelagic species underwent diurnal vertical migrations, while none of the bathypelagic species did so. The life histories of these pelagic fishes were compared among themselves and with epipelagic sardines and anchovies studied by others. The epipelagic species had the highest growth rates (estimated from otoliths, expressed in standard length or kilocalories), the mesopelagic species had the lowest growth rates and the bathypelagic species had intermediate growth rates. The relatively rapid growth rates of the bathypelagic fishes were achieved by high relative growth efficiencies made possible by low metabolic rates. Of the species studied, the lifespans of the epipelagic and bathypelagic species ranged from 4 to 8 yr and the lifespans of mesopelagic species from 5 to 8 yr. Data on egg diameters suggest that the mesopelagic species first reproduce in their 3rd yr, while the bathypelagic species do so in their last year. Epipelagic fishes generally have a large size, rapid growth, long life and early, repeated reproduction. Mesopelagic fishes are characterized by small size, slow growth, long life and early, repeated reproduction. Bathypelagic fishes generally have large size, rapid growth, somewhat shorter lives and late reproduction, which is possible a single event. The latter pattern is evidently feasible only in a rather stable environment where juvenile survivorship would always display relatively low variability. Many unusual characteristics of deep-living animals have possibly been selected by factors peculiar to the environment; however, such characteristics are just as likely to have been selected by factors equally present in many other environments, but not expressed there due to masking selective forces. In particular, we have in mind the darkness, stability and homogeneity of the bathypelagic realm as phenomena which represent the effective absence of many selective forces.The first two authors contributed equally and their names are listed in alphabetical order     

Gut length and mass in herbivorous and carnivorous prickleback fishes (Teleostei: Stichaeidae): ontogenetic, dietary, and phylogenetic effects

Relative gut length, Zihler’s index, and relative gut mass were measured in four species of prickleback fishes and the effects of ontogeny, diet, and phylogeny on these gut dimensions were determined. Of the four species, Cebidichthys violaceus and Xiphister mucosus shift to herbivory with growth (>45 mm SL), whereas X. atropurpureus and Anoplarchus purpurescens remain carnivores. A. purpurescens belongs to a carnivorous clade, and the three other species belong to an adjacent, herbivorous clade. Gut dimensions were compared in three feeding categories of the four species: (1) small, wild-caught juveniles representing the carnivorous condition before two species shift to herbivory; (2) larger, wild-caught juveniles representing the natural diet condition of the two carnivores and the two species that have shifted to herbivory; and (3) larger, laboratory-raised juveniles produced by feeding a high-protein artificial diet to small juveniles until they have reached the size of the larger, wild-caught juveniles. Comparisons of gut dimensions in categories (1) versus (2) tested for an ontogenetic effect, in (2) versus (3) for a dietary effect, and within each category for a phylogenetic effect. C. violaceus and X. mucosus increased gut dimensions with increase in body size and did not change ontogenetic trajectory in gut dimensions on the high-protein artificial diet, suggesting that they are genetically programmed to develop relatively large guts associated with herbivory. X. atropurpureus increased its gut dimensions with increase in size similar to its sister taxon, X. mucosus, suggesting a phylogenetic influence, but decreased gut dimensions on the high-protein artificial diet, suggesting phenotypic plasticity. Nevertheless, X. atropurpureus displayed a larger gut than A. purpurescens, further evidence that it evolved in an herbivorous clade. A. purpurescens possessed a relatively small gut that was little affected by ontogeny or diet. Ontogeny and phylogeny more than diet appear to influence gut dimensions in the four species, thus favoring genetic adaptation over phenotypic plasticity as the major force acting on digestive system features in the two prickleback clades.     

The anxious vole: the impact of group and gender on collective behavior under life-threat

Social animals behave collectively in order to maintain a cohesive group. This collective behavior is often led by a few individuals of specific gender, social rank, or spatial physical location in the group (i.e., perimeter or front). We examined how individual social voles (Microtus socialis) in same-gender compared with mixed-gender groups respond to an owl attack. We found that anxiety level, as measured by the time that each individual spent in less-sheltered sectors (open arms of elevated plus-maze and center of open arena), was affected by both the social context and the gender of the tested individuals. While both female and male voles generally reduced their activity in the open following owl attack, males in mixed-gender groups were exceptional in dichotomizing into those that spent a short period and those that spent a long period in the open arms of the plus-maze. Based on the similar responses of the same-gender groups, we suggest that anxiety is contagious, and based on the lower anxiety level of the mixed-gender groups, we suggest that natural groups that comprise both males and females are better able to cope with life-threat compared with same-gender groups. Finally, we suggest that the differential responses of males in the mixed-gender groups were due to a few males that displayed a low level of anxiety. These males were probably individuals of high social rank, and their response reflects their natural protective role, as previously described in social voles.