On the move around the clock: correlates and determinants of cathemeral activity in wild redfronted lemurs (<Emphasis Type="Italic">Eulemur fulvus rufus</Emphasis>)

Whether animals are active at night or during the day has profound consequences for many aspects of their behavioral ecology. Because of ecological and physiological trade-offs, most animals, including primates, are either strictly nocturnal or diurnal. However, a few primate species exhibit cathemeral activity, i.e., their activity is irregularly distributed throughout the 24-h cycle. Details and determinants of this unusual activity pattern are poorly understood because long-term 24-h observations are not feasible in the field. We therefore used small data loggers to record the activity of cathemeral redfronted lemurs (Eulemur fulvus rufus) from several neighboring groups quantitatively and continuously over a complete annual cycle in order to evaluate various proposed proximate and ultimate determinants of cathemeral activity. Activity data were examined for variation as a function of ambient temperature, time of day, lunar phase, and season. We found that cathemeral activity occurred year-round and that, on average, 3.5 times more activity occurred during the day. Total and diurnal activity increased during the long days of the austral summer. Nocturnal activity increased during the longer nights of the cool dry season. Irrespective of season, lunar phase had a significant effect on the distribution of activity across the 24-h cycle, with most nocturnal activity recorded during parts of the night with greatest brightness. These data indicate that light availability is the primary proximate determinant for the patterning of cathemeral activity. Several lines of evidence suggest that cathemerality in lemurs has evolved from nocturnal ancestors and that it represents a transitory state on the way to the diurnal niche.Communicated by F. Trillmich     

Feeding activity rhythm of<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> of the western Mediterranean shelf and slope grounds

Over a 24-h time scale, benthic decapods show complex patterns of behavioural activity associated with feeding. Information is scarce on the feeding rhythms of the burrowing decapod Nephrops norvegicus (L.), especially in those deep, benthic environments typical of western Mediterranean slopes, where the fishery of this species is highly developed. In the present study, the feeding of this species was determined by means of stomach sampling, from animals captured during four continuous 24-h cycles of trawl hauls at 100–110 m and 400–430 m, in October (the autumn equinox) and June (the spring solstice). A total of 3348 stomachs were analysed for fullness and classified by means of a visual, qualitative scale. A high proportion of the stomachs collected was empty (47.2%). Feeding rhythmicity was analysed by computing a percentage value of empty stomachs for each catch. Over 4 days, the percentage values presented marked fluctuations. Therefore, periodogram and form estimate analyses were performed on time series of percentages of empty stomachs, to determine the phase and periodicity of feeding rhythms. In October, animals ingested food during the daytime, presenting nocturnal peaks of empty stomachs at both depths considered in this study. In June, nocturnal peaks of empty stomachs were recorded for the slope, while a marked feeding arrhythmia was present on the shelf. A hypothesis linking feeding activity to endogenous rhythms in locomotion, cardiac and respiratory activities is presented to account for the animals diurnal ingestion, irrespective of the depth of distribution.Communicated by S.A. Poulet, Roscoff     

Influences of moonlight,ambient temperature,and food availability on the diurnal and nocturnal activity of owl monkeys (<Emphasis Type="Italic">Aotus azarai</Emphasis>)

The study of activity rhythms, their potential zeitgebers and masking factors among free-ranging primates has received relatively little attention in the past. Most primates are diurnal, a few of them nocturnal, and even fewer are cathemeral. Owl monkeys (Aotus azarai azarai) regularly show diurnal, as well as nocturnal, activity in the Argentinean and Paraguayan Chaco. The goal of this study was to examine how changes in activity patterns in owl monkeys of Formosa, Argentina are related to daily, monthly, and seasonal changes in temperature, light and food availability . During 1 year, I collected activity data from five groups followed continuously from dawn to dusk, dusk to dawn or uninterruptedly during 24 or 36 h for approximately 1,500 h. I kept hourly and daily records of temperature and light conditions, and I gathered monthly information on the density, distribution and abundance of food resources available to the monkeys. I found that the area of study is highly seasonal, and characterized by significant fluctuations in rainfall, temperature, photoperiod, and food availability. Owl monkeys had on average 5 h of activity during the day and 4 h during the night. The amount of diurnal activity remained fairly constant through the year despite seasonal changes in exogenous factors. Owl monkeys did not show changes in their activity patterns that could be attributed to changes in food availability. Nocturnal activity increased as the amount of moonlight increased, whereas diurnal activity decreased following a full-moon night. Ambient temperature was a good predictor of activity only when the moon was full. These results argue convincingly for an interaction between ambient temperature and moonlight in determining the observed activity pattern. It is then highly advisable that any evaluation of diurnal activity in cathemeral animals be analyzed controlling for the possible effects of moonlight during the previous night.Communicated by P. Kappeler     

Time stress,predation risk and diurnal–nocturnal foraging trade-offs in larval prey

Insect larvae increase in size with several orders of magnitude throughout development making them more conspicuous to visually hunting predators. This change in predation pressure is likely to impose selection on larval anti-predator behaviour and since the risk of detection is likely to decrease in darkness, the night may offer safer foraging opportunities to large individuals. However, forsaking day foraging reduces development rate and could be extra costly if prey are subjected to seasonal time stress. Here we test if size-dependent risk and time constraints on feeding affect the foraging–predation risk trade-off expressed by the use of the diurnal–nocturnal period. We exposed larvae of one seasonal and one non-seasonal butterfly to different levels of seasonal time stress and time for diurnal–nocturnal feeding by rearing them in two photoperiods. In both species, diurnal foraging ceased at large sizes while nocturnal foraging remained constant or increased, thus larvae showed ontogenetic shifts in behaviour. Short night lengths forced small individuals to take higher risks and forage more during daytime, postponing the shift to strict night foraging to later on in development. In the non-seasonal species, seasonal time stress had a small effect on development and the diurnal–nocturnal foraging mode. In contrast, in the seasonal species, time for pupation and the timing of the foraging shift were strongly affected. We argue that a large part of the observed variation in larval diurnal–nocturnal activity and resulting growth rates is explained by changes in the cost/benefit ratio of foraging mediated by size-dependent predation and time stress.     

Arrived in the light: diel and seasonal activity patterns in wild Verreaux’s sifakas (<Emphasis Type="Italic">Propithecus v. verreauxi</Emphasis>; Primates: Indriidae)

Madagascar is characterised by pronounced annual climatic and ecological seasonality and harbours a radiation of closely related sympatric primates (Lemuriformes) that exhibit diurnal, nocturnal and cathemeral activity patterns. We collected continuous activity data over a complete annual cycle from wild diurnal Verreauxs sifakas (Propithecus v. verreauxi) to contribute detailed and comparative data: (1) to characterise their diel and seasonal activity rhythms, (2) to identify factors shaping variation in activity rhythms, and (3) to help reconstruct the evolutionary transition from nocturnal to diurnal activity. We fitted eight adult sifakas from seven different groups living in Kirindy forest with an accelerometer/data logger device and recorded their activity in 5-min bins for a total of 12 months. We characterise P. verreauxi as a strictly diurnal species with a pronounced bimodal activity pattern that briefly changed to a more unimodal pattern during their annual mating season (January to March). We documented significant annual variation in total daily activity, activity time, and activity level, as well as in most parameters characterising their bimodal activity pattern. Despite a significant positive correlation of the animals activity time with day length, pronounced annual variation in the phase position of onset and end of activity in relation to sunrise and sunset times could also be discerned. Minor enhancing effects of moonlight on nocturnal activity were only found for the first 3 h of the inactivity period. Bimodality of the activity pattern and the additional reduction of activity time during the cold and dry winter months associated with reduced food availability can be interpreted as flexible behavioural adaptations to reduce energy expenditure. We therefore propose that energetic and thermoregulatory benefits are important factors shaping these primates activity pattern.Communicated by C. Nunn     

Evaluating day–night changes in shallow Mediterranean rocky reef fish assemblages by visual census

Ecological information on coastal fish distribution patterns and habitat use can be greatly improved by nocturnal samplings and observations. To this purpose, the structure of a Mediterranean fish assemblage inhabiting the shallow rocky littoral of Linosa Island (Sicily Strait, Italy) was examined by using visual census to detect possible diel variations in species composition and abundance. Day–night fish distribution patterns were investigated by multivariate and univariate analyses. Overall, 42 fish taxa belonging to 19 families were recorded: 35 during the day and 24 during the night. Seventeen species were common to both diurnal and nocturnal assemblages. Within the diurnal assemblage, Chromis chromis was the most represented species (37.2%), followed by Thalassoma pavo (23.2%) and Sparisoma cretense (10.8%). Within the nocturnal assemblage, the most abundant taxon was Atherina spp. (33.9%), followed by Apogon imberbis (26.4%) and Boops boops (11.5%). Our results indicated wide variation in the abundance and species composition during the day and during the night. Multi Dimensional Scaling plot showed a clear-cut separation between the two assemblages and analysis of similarities found significant differences as well. SIMPER analysis revealed that ten species individually contributed by more than 2.5% to the dissimilarity between diurnal and nocturnal assemblages, T. pavo, C. chromis and S. cretense being the first three species in order of decreasing percentage. ANOVA performed on species richness and fish abundance detected significant differences between diurnal and nocturnal assemblages, the latter showing far lower average values for both variables.     

Lemurs and the Regeneration of Dry Deciduous Forest in Madagascar

Abstract: We sought to assess the role of lemurs for seed dispersal in the dry deciduous forest of western Madagascar and the possible consequences of the demise of lemurs for forest regeneration. Forest regeneration was studied in eight plots in two large blocks of primary forest and in seven fragments of primary forest (1 plot per fragment). In 4 of the 15 study plots, the abundance of saplings was negatively and significantly correlated ( p < 0.05) with the abundance of mature individuals of the same tree species. In another 10 study plots there were negative correlations, although these were not significant on the community level. Second-order statistics were significant with p < 0.001 and indicated that seed dispersal away from the parent trees was important for successful establishment of saplings. Apart from possibly the bush pig ( Potamochoerus larvatus ), only one vertebrate species of the dry forest, the brown lemur ( Eulemur fulvus ), ingested seeds> 11 mm long and passed them through its digestive tract unharmed. These results for lemurs were based on direct observations and fecal analyses. To evaluate the role of E. fulvus , we compared regeneration in forest plots with and without E. fulvus . In forest fragments without E. fulvus , fewer lemur-dispersed tree species regenerated than would be expected based on the presence of mature tree species that are lemur-dispersed ( p < 0.05). No such effect was seen in primary forests with E. fulvus or for trees whose seeds can also be dispersed by other vertebrates. Thus, regeneration of the dry deciduous forest of western Madagascar with the complete set of primary forest tree species seems to depend upon the presence of E. fulvus .     

Mutual tolerance or reproductive competition? Patterns of reproductive skew among male redfronted lemurs (<Emphasis Type="Italic">Eulemur fulvus rufus</Emphasis>)

The social organization of gregarious lemurs significantly deviates from predictions of the socioecological model, as they form small groups in which the number of males approximately equals the number of females. This study uses models of reproductive skew theory as a new approach to explain this unusual group composition, in particular the high number of males, in a representative of these lemurs, the redfronted lemur (Eulemur fulvus rufus). We tested two central predictions of “concession” models of reproductive skew theory, which assume that subordinates may be allowed limited reproduction by dominant group members as an incentive to remain in the group, thereby increasing the group’s overall productivity. Accordingly, relatives are predicted to receive less reproduction than non-relatives, and the overall amount of reproductive concessions given to subordinates is predicted to increase as the number of subordinates increases. In addition, we tested whether the number of females in a group, a variable not previously incorporated in reproductive skew theory, affected reproductive skew among males. Using microsatellite analyses of tissue DNA, we determined paternities of 49 offspring born into our study population in Kirindy forest (western Madagascar) since 1996 to determine patterns of male reproductive skew to test these predictions. Our analyses revealed remarkable reproductive skew, with 71% of all infants being sired by dominant males, but both predictions of reproductive skew models could not be supported. Instead, the number of females best predicted the apportionment of reproduction among the males in this species, suggesting that current reproductive skew models need to incorporate this factor to predict reproductive partitioning among male primates and perhaps other group-living mammals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Both Peter M. Kappeler and Markus Port contributed equally to this paper.     

Population viability and harvest sustainability for Madagascar lemurs

Subsistence hunting presents a conservation challenge by which biodiversity preservation must be balanced with safeguarding of human livelihoods. Globally, subsistence hunting threatens primate populations, including Madagascar's endemic lemurs. We used population viability analysis to assess the sustainability of lemur hunting in Makira Natural Park, Madagascar. We identified trends in seasonal hunting of 11 Makira lemur species from household interview data, estimated local lemur densities in populations adjacent to focal villages via transect surveys, and quantified extinction vulnerability for these populations based on species-specific demographic parameters and empirically derived hunting rates. We compared stage-based Lefkovitch with periodic Leslie matrices to evaluate the impact of regional dispersal on persistence trajectories and explored the consequences of perturbations to the timing of peak hunting relative to the lemur birth pulse, under assumptions of density-dependent reproductive compensation. Lemur hunting peaked during the fruit-abundant wet season (March–June). Estimated local lemur densities were roughly inverse to body size across our study area. Life-history modeling indicated that hunting most severely threatened the species with the largest bodies (i.e., Hapalemur occidentalis, Avahi laniger, Daubentonia madagascariensis, and Indri indi), characterized by late-age reproductive onsets and long interbirth intervals. In model simulations, lemur dispersal within a regional metapopulation buffered extinction threats when a majority of local sites supported growth rates above the replacement level but drove regional extirpations when most local sites were overharvested. Hunt simulations were most detrimental when timed to overlap lemur births (a reality for D. madagascariensis and I. indri). In sum, Makira lemurs were overharvested. Regional extirpations, which may contribute to broad-scale extinctions, will be likely if current hunting rates persist. Cessation of anthropogenic lemur harvest is a conservation priority, and development programs are needed to help communities switch from wildlife consumption to domestic protein alternatives.     

The lemur syndrome unresolved: extreme male reproductive skew in sifakas (<Emphasis Type="Italic">Propithecus verreauxi</Emphasis>), a sexually monomorphic primate with female dominance

The primates of Madagascar (Lemuriformes) are unusual among mammals in that polygynous species lack sexual dimorphism, and females dominate males socially in most species. Moreover, lemur groups are relatively small and characterized by even adult sex ratios despite the fact that one male should be able to exclude other males from the group. One hypothesis to explain this combination of behavioral, morphological, and demographic traits (the “lemur syndrome”) postulates that male–male competition is relaxed and, hence, variance in male reproductive success is low. Reproductive skew theory provides a framework for testing this and several related predictions about lemur social evolution. Specifically, low reproductive skew is also predicted if dominant males or adult females make reproductive concessions to subordinates or if the latter group successfully pursues alternative reproductive tactics. However, suitable data on paternity, demography, and behavior for a conclusive test of these predictions have not been available in the past. In this paper, we show that male reproductive success in ten groups of Verreaux’s sifakas (Propithecus verreauxi) was extremely skewed in favor of the dominant male over 9 years. Our genetic analyses also revealed that more than a third of all groups are effectively harem groups because only one male was unrelated to the resident female(s). In groups with two or more non-natal males, the dominant sired 91% of 33 infants. Together, males pursuing one of several alternative reproductive tactics, such as roaming among several groups or immigrating peacefully, sired only 11% of infants. Thus, female sifakas do not control group composition by offering reproductive opportunities to subordinate males as staying incentives, intrasexual selection is not relaxed, and dominant males prevail in a tug-of-war over subordinate males. Because male reproductive skew in sifakas is even more pronounced than in harem-living anthropoids studied to date, intrasexual selection is clearly not relaxed, and the lemur syndrome is more puzzling than ever.     

To breed or not to breed—food competition and other factors involved in female breeding decisions in the pair-living nocturnal fork-marked lemur (<Emphasis Type="Italic">Phaner furcifer</Emphasis>)

Differential access to food resources is thought to be the main determinant of differences in female reproductive success but is poorly studied in both pair-living and nocturnal primates. The modes of food competition within and between families were investigated following the principles proposed by the ecological model using 3 years of field data from seven fork-marked lemur (Phaner furcifer) families. The major food resources were identified from year-round feeding observations and the strength and mode of competition were inferred from differences in physical condition. The most important food resource of fork-marked lemurs were tree exudates which occurred in small, defendable food patches, characterized by fast depletion and rapid renewal. These characteristics led to strong within-group contest and scramble competition, which were found to yield a positive dominance effect and a negative group-size effect on female net energy gain. Differential physical condition, however, did not translate directly into differential reproductive success. Low female fertility was best predicted by large family size associated with delayed dispersal by previous offspring. Although there is no obvious benefit from full-grown offspring in their territory, adults tolerate delayed natal dispersal, probably because dispersal poses extraordinary costs for the offspring. These costs are likely to accrue from decreased foraging efficiency in unfamiliar habitats because exudate feeding requires very rigid feeding itineraries. In conclusion, the presented evidence for group-size effects on reproductive success in pair-living females opens a new area for research on the costs and benefits of delayed dispersal and female reproductive decisions.This revised version was published in September 2003 with corrections to the Authors Present address.An erratum to this article can be found at Communicated by E.H.M. Sterck     

Transfer of brevetoxins to a tellinid bivalve by suspension- and deposit-feeding and its implications for clay mitigation of <Emphasis Type="Italic">Karenia brevis</Emphasis> blooms

Blooms of the brevetoxin-producing Karenia brevis in the Gulf of Mexico cause massive fish kills, food poisoning and adverse respiratory effects in humans. Sedimentation of toxic cells following inert clay application could reduce toxin incorporation by commercially important suspension-feeding bivalves and thus prevent direct public health impacts, but could potentially lead to brevetoxin (PbTx) accumulation by benthic deposit-feeders. The goal of this study was therefore to compare suspension- and deposit-feeding as pathways for brevetoxins. We investigated: (1) the effect of toxic K. brevis on both feeding modes using a facultative deposit-suspension feeding tellinid bivalve, the clam Macoma balthica, as a model species and (2) the relative effectiveness of brevetoxin transfer via suspension- and deposit-feeding over 24-h exposure. Sedimentation of K. brevis was achieved by treatment with 0.25 g phosphatic clay l⁻¹ and brevetoxin concentrations were measured by ELISA. Karenia brevis reduced both suspension- and deposit-feeding activity. This study demonstrates that brevetoxins can be rapidly accumulated by a surface deposit-feeding bivalve from sedimented K. brevis cells and that comparable toxin levels can be attained by both suspension- and deposit-feeding modes [1.2–1.6 μg PbTx (g tissue wet weight)⁻¹]. Deposit-feeding clams generally do not pose a direct threat to humans but may provide a pathway for brevetoxin food web transfer.     

Small testes size despite high extra-pair paternity in the pair-living nocturnal primate Phaner furcifer

Sperm competition will be the inevitable consequence of polyandrous mating behavior if two or more males inseminate a single female. It has been demonstrated for a wide variety of animals that males adapt to this situation behaviorally, physiologically and morphologically, e.g. by evolving relatively large testes size to produce more sperm. All pair-living primates investigated so far were found to have relatively small testes, suggesting a monandrous mating system. We investigated the relationship between extra-pair paternity (EPP) rate as a measure of sperm competition intensity and relative testes size in a pair-living primate, the fork-marked lemur (Phaner furcifer). Paternity exclusion analyses for seven offspring using six polymorphic DNA-microsatellite markers suggested a high EPP rate. Female nocturnal travel distances were longer during the mating season, suggesting that females take an active role in achieving extra-pair copulations (EPCs). Surprisingly, fork-marked lemur testes size was relatively small compared to 23 other lemuroid primates, a result that is in contrast to predictions of sperm competition theory. Neither possible behavioral and morphological adaptations to an alternative paternity guard (i.e. mate guarding), nor sampling biases, phylogenetic constraints, and population density effects explain the absence of large testes in a species with high EPP, a phenomenon also known from birds with moderate to low EPP rates. We conclude that more data are needed on the frequency of EPCs, the timing of in-pair and extra-pair copulations, as well as the role of female choice, to explain why males of some species apparently do not adapt to sperm competition.Communicated by S. Alberts     

Seasonal variation in the effects of food availability on gametogenesis in the purple urchin (<Emphasis Type="Italic">Strongylocentrotus purpuratus</Emphasis>)

The amount of food and when it is available affect both the timing of reproduction and the number of gametes produced by purple sea urchins, Strongylocentrotus purpuratus. To investigate this further, the effects of food availability on feeding rates, gonad growth, and gamete development were examined in S. purpuratus collected from the Point Loma kelp forest near San Diego, California, USA (32.69° N, 117.26° W) in September (Fall) 2007, and February (Spring) and July (Summer) 2008, using laboratory mesocosms. Each seasonal laboratory feeding experiment lasted 3 months, and different levels of food availability were established with different feeding frequencies (from 1 to 7 days week⁻¹). Gonad tissues of male and female urchins were staged at the end of each experiment using histological analyses. Reduced food availability resulted in increased daily consumption rates, especially in the Fall when gamete development began. Food limitation at this time resulted in failure to produce viable gametes, suggesting there is a critical period early in gonad development when food limitation affects reproductive competency. Food limitation later in gonad development did not stop viable gamete production, although it did reduce gamete output.     

Relative intensity of predation on the French grunt, Haemulon flavolineatum, during diurnal, dusk, and nocturnal periods on a coral reef

Established hypotheses state that the rate of predation on coral reef fish should be highest during crepuscular periods (dawn and dusk) intermediate diurnally, and lowest nocturnally. We examined the relative risk of predation on juvenile French grunts (Haemulon flavolineatum Desmarest) during diurnal, dusk, and nocturnal periods on the fore- and back-reef at Teague Bay, U.S. Virgin Islands in July and August 1996. Tethering-devices recorded the exact amount of time between attaching a prey fish to its tether and subsequent predation on the prey fish. As tethering of prey usually inflates the actual rate of predation, times from our tethering devices were used to establish only the relative predation risk among treatments. During 3-h diurnal and nocturnal tethering experiments, relative predation was significantly higher during the nocturnal period, and differences between side of reef were not significant. In 30-min tethering experiments, which included all three time periods, the relative predation risk was significantly higher during dusk and nocturnal periods than during the diurnal period. Relative predation was not significantly different between the dusk and nocturnal periods, or between side of reef during any time period. The unexpected finding that the diurnal period had the lowest relative risk of predation indicates that the timing of predation events on reefs, as well as the adaptive reasons for nocturnal larval settlement, may need to be re-examined. Received: 11 February 1997 / Accepted: 21 October 1998     

Behavioural responses of the Dungeness crab, <Emphasis Type="Italic">Cancer magister</Emphasis>, during feeding and digestion in hypoxic conditions

The Dungeness crab, Cancer magister, inhabits areas that are frequently subject to periods of hypoxia. This species can employ physiological mechanisms that allow it to cope with acute hypoxic episodes. When crabs feed there is a general increase in physiological variables; these may pose an additional physiological burden on crabs already attempting to maintain adequate oxygen uptake in hypoxia. In Barkley Sound, British Columbia, the inshore habitats of C. magister ranged in dissolved oxygen from 28 kPa at the water surface to less than 1.0 kPa just above the sediment–water interface. During short-term hypoxic events, crabs reduced both the amount of food eaten and the amount of time spent feeding. Crabs tended to cease feeding below 3.2 kPa oxygen, but resumed feeding when the dissolved oxygen tensions were rapidly raised to 6 kPa. In a high (10.5–21 kPa) oxygen gradient, both unfed and fed crabs showed no preference for any area of the gradient. In a low (2.5–10.5 kPa) dissolved oxygen gradient, both unfed and fed crabs preferred the highest oxygen regime. In the laboratory, crabs were less likely to enter hypoxic waters (below 3.2 kPa oxygen) to obtain and consume food; those that did moved the food to a higher oxygen regime prior to feeding and settled in higher oxygen regimes for digestion. Crab behaviour was also monitored in the field. Fed and unfed crabs were fitted with ultrasonic telemetry tags and tracked during a tidal cycle. Unfed crabs remained mobile, travelling up to 1,370 m within 6 h, while postprandial crabs settled in areas of high oxygen and moved very little during the first 48 h after release. The present study suggests that C. magister exhibits behavioural responses in order to minimise the use of physiological mechanisms, and maximise foraging and digestive processes. Thus the nutritional state of the individual may be important in regulating both its behaviour and distribution in its natural environment.     

Androgen and glucocorticoid levels reflect seasonally occurring social challenges in male redfronted lemurs (Eulemur fulvus rufus)

Intense reproductive competition and social instability are assumed to increase concentrations of glucocorticoids and androgens in vertebrates, as a means of coping with these challenges. In seasonally breeding redfronted lemurs (Eulemur fulvus rufus), the mating and the birth season and the associated increased male competition are predicted to pose such reproductive challenges. In this paper, we investigate seasonal variation in hormone excretion in male redfronted lemurs, and examine whether this variation is associated with social or ecological factors. Although dominance status has been shown to affect individual stress levels across many taxa, we predicted no rank-related differences in glucocorticoids for redfronted lemurs because relatively equal costs are associated with both high and low rank positions (based on patterns of rank acquisition/maintenance and threats toward subordinates). Over a 14-month period, we collected behavioral data (1843 focal hours) and 617 fecal samples from 13 redfronted lemur males in Kirindy Forest/Madagascar. We found no general rank-related pattern of testosterone or glucocorticoid excretion in this species. Both hormones were excreted at significantly higher levels during the mating and the birth season, despite social stability during both periods. The elevated mating season levels may be explained by increased within-group reproductive competition during this time and are in line with previous studies of other seasonally reproducing primates. For the birth season increase, we propose that the predictable risk of infanticide in this highly seasonal species affects male gonadal and adrenal endocrine activity. We evaluate alternative social and ecological factors influencing the production of both hormone classes and conclude based on our preliminary investigations that none of them can account for the observed pattern.     

A dusk chorus effect in a nocturnal bird: support for mate and rival assessment functions

The dawn chorus is a striking feature of spring mornings and a characteristic behaviour of many bird species, particularly the passerines. Dawn singing has been considered a reliable signal of male quality for mate and rival assessment. Singing is presumed to be relatively costly at dawn both because air temperatures are relatively low and because birds have not fed overnight. Models of optimal daily routine predict the existence of a “dusk chorus” in nocturnal birds, although this prediction has received little empirical attention. Nocturnal birds at dusk may be energy-limited because of a lack of daytime feeding, and singing at dusk may thus ensure signal reliability. Here, we used an observational and experimental approach to study vocal behaviour at dusk and dawn in a nocturnal raptor, the little owl Athene noctua. We assess whether male little owls adjust their vocal behaviour according to feeding stage (i.e. period of the night), ambient air temperature and territorial context (i.e. spontaneous calling behaviour vs elicited calling by intrusion). Across different temperatures, we find that both spontaneous vocal activity and inter-individual variability in call duration increased at dusk, clearly indicating a dusk chorus phenomenon. Results from playback presentations suggest that food, rather than air temperature, is likely to be more constraining at dusk. We discuss how comparing dusk and dawn choruses in nocturnal and diurnal species can provide insights into both mechanistic and functional aspects of signalling behaviour.     

<Emphasis Type="Italic">Octopus insularis</Emphasis> (Octopodidae), evidences of a specialized predator and a time-minimizing hunter

Shallow-water octopuses have been reported as major predators of motile species in benthonic marine communities, capturing their prey by different foraging techniques. This study assessed for the first time the feeding ecology, foraging behavior, and defensive strategy during foraging, including the use of body patterns, to construct a general octopus foraging strategy in a shallow water-reef system. Octopus insularis was studied in situ using visual observations and video recordings. The diet included at least 55 species of crustaceans (70%), bivalves (17.5%), and gastropods (12.5%); however, only four species accounted for half of the occurrences: the small crabs Pitho sp. (26.8%) and Mithrax forceps (23.9%), the bivalve Lima lima (5.3%), and the gastropod Pisania pusio (4.9%). Poke and crawl were most frequent foraging behaviors observed in the video recordings. The foraging behaviors were associated with environmental variables and octopus body size. The sequences of foraging behavior showed characteristics of a tactile saltatory searching predator, as well as a visual opportunist. Body patterns showed a relationship with foraging behavior, habitat variables, and octopus body size. Mottle was the most frequent pattern, especially during poke and crawl, in shallower depths. Dorsal light–ventral blue green was more frequent during swimming at mid-water, and Blotch was the normal pattern during web-over by large animals. The large proportion of two species of small crabs in den remains, the intense search for food during short hunting trips, and the intense use of cryptic body patterns during foraging trips, suggest that this species is a ‘time-minimizing’ forager instead of a ‘rate-maximizer’.     

Communal nesting, kinship, and maternal success in a social primate

Communal nesting, where several mothers regularly pool and cooperatively rear offspring, is unusual in mammals. This type of crèching behavior is especially rare among primates, with the notable exceptions of humans, some nocturnal strepsirrhines, and—as we show in this study—black-and-white ruffed lemurs (Varecia variegata). Here, we combine data on nesting behavior, genetic relatedness, and infant survivorship to describe variation in ruffed lemur infant care and to examine the potential benefits of ruffed lemur communal breeding. Reproductive events were rare, and females produced litters (synchronously) only once in 6 years of observation. We show that not all mothers participate in communal crèches, but those that did had greater maternal success; communal breeders spent more time feeding and their offspring were more likely to survive. Although cooperating mothers were often related, females also cooperated with non-kin, and those who shared infant care responsibilities had greater maternal success than mothers who did not participate. If there is indeed a causal link between maternal cooperation and reproductive success, this unusual behavior, like that of human communal rearing, may have evolved via some combination of kin selection and mutualism.