Sociality influences thermoregulation and roost switching in a forest bat using ephemeral roosts

In summer, many temperate bat species use daytime torpor, but breeding females do so less to avoid interferences with reproduction. In forest‐roosting bats, deep tree cavities buffer roost microclimate from abrupt temperature oscillations and facilitate thermoregulation. Forest bats also switch roosts frequently, so thermally suitable cavities may be limiting. We tested how barbastelle bats (Barbastella barbastellus), often roosting beneath flaking bark in snags, may thermoregulate successfully despite the unstable microclimate of their preferred cavities. We assessed thermoregulation patterns of bats roosting in trees in a beech forest of central Italy. Although all bats used torpor, females were more often normothermic. Cavities were poorly insulated, but social thermoregulation probably overcomes this problem. A model incorporating the presence of roost mates and group size explained thermoregulation patterns better than others based, respectively, on the location and structural characteristics of tree roosts and cavities, weather, or sex, reproductive or body condition. Homeothermy was recorded for all subjects, including nonreproductive females: This probably ensures availability of a warm roosting environment for nonvolant juveniles. Homeothermy may also represent a lifesaver for bats roosting beneath loose bark, very exposed to predators, because homeothermic bats may react quickly in case of emergency. We also found that barbastelle bats maintain group cohesion when switching roosts: This may accelerate roost occupation at the end of a night, quickly securing a stable microclimate in the newly occupied cavity. Overall, both thermoregulation and roost‐switching patterns were satisfactorily explained as adaptations to a structurally and thermally labile roosting environment.     

Roosting Behavior of Colonial and Solitary Flying Foxes in American Samoa (Chiroptera: Pteropodidae)1

We examined characteristics of roosting sites utilized by two flying fox species (Pteropus tonganus and P. samoensis) in American Samoa. The colonial roosting sites of P. tonganus were observed over a ten‐year period, including two years when severe hurricanes devastated bat populations and destroyed roost trees. Prior to the hurricanes, roosts were located on cliff faces above the ocean or steep mountainsides, locations that were either inaccessible to people or in protected areas where hunting was not allowed. In the years immediately following the hurricanes, P. tonganus colonies split into smaller groups that moved frequently to different locations. Four years after the second hurricane, colonies had coalesced and returned to many of the traditional roosting sites used before the hurricanes. Common tree species in upland and coastal forest were selected as roosts. The isolated locations selected for P. tonganus roosts were apparently the result of hunting pressure on the colonies. The solitary roosts of P. samoensis were observed during 29 months. Roosting bats were well concealed and hard to detect within the forest; even bats on exposed branches were cryptic. Mature primary forest was favored as roosting habitat. Individual bats used specific branches or trees as roosts and returned to them for up to 29 months. Unlike P. tonganus, people did not alarm roosting P. samoensis easily and some roosts were located near houses and along roads.     

Restless roosts: Light pollution affects behavior,sleep, and physiology in a free‐living songbird

The natural nighttime environment is increasingly polluted by artificial light. Several studies have linked artificial light at night to negative impacts on human health. In free‐living animals, light pollution is associated with changes in circadian, reproductive, and social behavior, but whether these animals also suffer from physiologic costs remains unknown. To fill this gap, we made use of a unique network of field sites which are either completely unlit (control), or are artificially illuminated with white, green, or red light. We monitored nighttime activity of adult great tits, Parus major, and related this activity to within‐individual changes in physiologic indices. Because altered nighttime activity as a result of light pollution may affect health and well‐being, we measured oxalic acid concentrations as a biomarker for sleep restriction, acute phase protein concentrations and malaria infection as indices of immune function, and telomere lengths as an overall measure of metabolic costs. Compared to other treatments, individuals roosting in the white light were much more active at night. In these individuals, oxalic acid decreased over the course of the study. We also found that individuals roosting in the white light treatment had a higher probability of malaria infection. Our results indicate that white light at night increases nighttime activity levels and sleep debt and affects disease dynamics in a free‐living songbird. Our study offers the first evidence of detrimental effects of light pollution on the health of free‐ranging wild animals.     

The Red Fox,Vulpes vulpes,as a Predator upon Earthworms,Lumbricus terrestris

The tiny (3.1–3.8 g) vespcrtilionid bat Pipistrellus nanus was studied in Kenya palm-thatched roofs from May 1973 to July 1974. Roosting social organization and related activities and behavior are described. ♂♂ held diurnal roosting territories where ♀♀ gathered in small and compositionally labile groups, attracted to the most vocal ♂♂. Annual variation in population-wide aspects of social organization follows predictable seasonal changes in climate and predator abundance. Variability between individuals follows a common mammalian pattern: high male competition for ♀, variance in presumed male reproductive success, and a mating system resembling one based on resource defense polygyny. Social organization in this population contrasts with that known from studies of other P. nanus populations.     

Night roosting and the nocturnal time budget of the little brown bat,Myotis lucifugus: Effects of reproductive status,prey density,and environmental conditions

Summary The insectivorous bat Myotis lucifugus typically apportions the night into two foraging periods separated by an interval of night roosting. During this interval, many bats occupy roosts that are used exclusively at night and are spatially separate from maternity roosts. The proportion of the night which bats spend roosting, and thus the proportion spent foraging, vary both daily and seasonally in relation to the reproductive condition of the bats, prey density, and ambient temperature. A single, continuous night roosting period is observed during pregnancy. During lactation, females return to maternity roosts between foraging bouts, and night roosts are used only briefly and sporadically. Maximum use of night roosts occurs in late summer after young become volant. Superimposed upon these seasonal trends is day-to-day variation in the bats' nightly time budget. Long night roosting periods and short foraging periods are associated with cool nights and low prey density. This behavioral response may minimize energetic losses during periods of food scarcity.     

Genetic mating system and the significance of harem associations in the bat Saccopteryx bilineata

We analysed the polygynous mating system of the bat Saccopteryx bilineata using behaviour observations and genetic data on 11 microsatellite DNA loci. Basic social units in S. bilineata are harem groups that consist of single males and up to eight females. Colonies comprise several harem groups, and the composition of colonies and harems is often stable over several reproductive seasons. The combination of parentage exclusion and likelihood-based parentage assignment in this study produced detailed parentage information for a large colony of S. bilineata. Reproduction occurred mostly within the colony (17% extra-colony paternity), but social associations in harems within the colony did not represent reproductive units (70% extra-harem paternity). The latter finding was consistent over three reproductive seasons. Spatial association of the roosting sites of males and females could not explain parentage patterns in the colony. Even though intra-harem paternity was less frequent than expected, it contributed significantly to reproduction of harem males. On average, the number of offspring sired by a male with females in his harem territory increased significantly with harem size, which corresponds to the higher energetic investment that is related to the maintenance of large harems. However, extra-harem paternity was not correlated with a male's harem size or intra-harem reproductive success. This suggests that individual preferences of females rather than male traits associated with the ability to defend large harems are most likely to cause the detected differences between social association and genetic mating system.     

云南蒙自钳嘴鹳夜栖地利用及夜栖树选择

动物对夜栖地选择的灵活程度是其分布和扩散的重要因素之一。钳嘴鹳(Anastomus oscitans)为中国新分布种,要了解其扩散趋势,掌握其夜栖地需求至关重要。2015年11月至2018年7月在云南蒙自坝区,使用卫星跟踪技术对6只钳嘴鹳进行了夜栖地利用和夜栖树选择研究,结果表明:(1)钳嘴鹳可以在多种生境中夜栖,包括林地、沼泽、岛屿等。从群体水平上分析,钳嘴鹳主要夜栖于沼泽中,利用率为(51.19±12.34)%,其次为林地(31.55±11.34)%和岛屿(17.26±5.70)%;从个体水平上来看,不同钳嘴鹳的夜栖地利用方式差异较大,其中4只主要利用沼泽,而另外2只主要利用林地;(2)随机森林分析表明:影响钳嘴鹳夜栖树选择的首要因子为距觅食地距离,其次为树高、地径和最低枝高度,而人为干扰因子的影响较弱。综合来看,钳嘴鹳对夜栖地类型以及夜栖树均表现出了极强的适应性和可塑性,这可能是其能够快速扩散的重要原因之一。同时也表明夜栖树并不能成为一个有力的限制因子,钳嘴鹳有可能继续向国内的南方湿地区扩散。     

Roosting Ecology of the Tent‐Roosting Bat Artibeus watsoni (Chiroptera: Phyllostomidae) in Southwestern Costa Rica1

We described the plants used as roost resources by Artibeus watsoni in southwestern Costa Rica, assessed roost fidelity, and compared roosting ecology between two sites, Golfito and Corcovado, which vary in the degree of human influence. A total of 349 tents from 25 different plant species were used by A. watsoni as roosts; some plant species (e.g., Carludovica palmata, Asplundia alata, Heliconia imbricata and Calathea lutea) were modified into tents with significantly higher frequency than others. The highest tents above the ground were observed in Philodendron popenoei and Rhodospatha wendlandii, whereas tents in Philodendron grandipes and A. alata were significantly lower than any other species. Asplundia alata and R. wendlandii also had the highest frequency of leaves modified per plant. Fidelity of bats to tents was low, although bats used several tents intermittently within a restricted area. Males generally were more faithful to tents than females, although not significantly so. This observation, along with indirect evidence of leaf modification, suggests that males are primarily responsible for tent construction. The two study sites differed in the plants used for roosting and in tent fidelity. Bats in Corcovado used a greater variety of plant species for tent roosting, whereas bats in Golfito were more faithful, suggesting that roosting resources were scarcer at the latter site.     

Social calls used by a leaf-roosting bat to signal location

Social calls in bats have many functions, including mate attraction and maintaining contact during flight. Research suggests that social calls may also be used to transfer information about roosts, but no studies have yet demonstrated that calls are used to actively attract conspecifics to roosting locations. We document the social calls used by Spix''s disc-winged bat (Thyroptera tricolor) to actively recruit group members to roosts. In acoustic trials, we recorded two sets of calls; one from flying individuals termed ‘inquiry calls’, and another from roosting bats termed ‘response calls’. Inquiry calls were emitted by flying bats immediately upon release, and quickly (i.e. 178 ms) elicited production of response calls from roosting individuals. Most flying bats entered the roost when roosting individuals responded, while few bats entered the roost in the absence of a response. We argue that information transfer concerning roost location may facilitate sociality in T. tricolor, given the ephemeral nature of roosting structures used by this species.     

Reproductive system,social organization,human disturbance and ecological dominance in native populations of the little fire ant,Wasmannia auropunctata

The invasive ant species Wasmannia auropunctata displays both ecologically dominant and non‐dominant populations within its native range. Three factors could theoretically explain the ecological dominance of some native populations of W. auropunctata: (i) its clonal reproductive system, through demographic and/or adaptive advantages; (ii) its unicolonial social organization, through lower intraspecific and efficient interspecific competition; (iii) the human disturbance of its native range, through the modification of biotic and abiotic environmental conditions. We used microsatellite markers and behavioural tests to uncover the reproductive modes and social organization of dominant and non‐dominant native populations in natural and human‐modified habitats. Microsatellite and mtDNA data indicated that dominant and non‐dominant native populations (supercolonies as determined by aggression tests) of W. auropunctata did not belong to different evolutionary units. We found that the reproductive system and the social organization are neither necessary nor sufficient to explain W. auropunctata ecological dominance. Dominance rather seems to be set off by unknown ecological factors altered by human activities, as all dominant populations were recorded in human‐modified habitats. The clonal reproductive system found in some populations of W. auropunctata may however indirectly contribute to its ecological dominance by allowing the species to expand its environmental niche, through the fixation over time of specific combinations of divergent male and female genotypes. Unicoloniality may rather promote the range expansion of already dominant populations than actually trigger ecological dominance. The W. auropunctata model illustrates the strong impact of human disturbance on species’ ecological features and the adaptive potential of clonal reproductive systems.     

Analysing hierarchically-structured fitness and modular dynamics in plants: integration of concepts from population dynamics

This paper analyses the reproductive strategies of a modular organism by means of demographic analysis of its reproductive units. The buds of the reproductive structure of Retama sphaerocarpa (L.) Boiss., a perennial shrub with a simple modular structure, were considered as ‘individuals’ of a population of modules within each individual plant. The development of the buds leads to the production of subpopulations of new units (inflorescences, flowers, fruits) of a lower hierarchical level. Fitness and hierarchical fitness (defined by the integration of developmental hierarchical levels in the plant) were analysed at the shoot module level, starting from the analogy provided by classical population dynamics and taking into account the different subpopulations of modules that take part in the reproductive process. Consideration of the dynamics of such subpopulations reveals the way in which demographic strategies partitioning the reproductive effort are exerted throughout the plant, and their consequences for fitness. The analysis of the reproductive process in R. sphaerocarpa shows a critical developmental transition from flowering to fruiting buds, which is a consequence of a low survival rate of the subpopulation of flowers. Despite the specificity of the empirical information used in this investigation, the fitness analyses proposed in this paper are fully applicable to any hierarchically-structured biological system, either modular or unitary.     

Tests of hypotheses for group formation in the subtropical leaf‐dwelling bat,Kerivoula furva

Investigating factors that promote group living in animals can help us to understand the evolution of sociality. The dark woolly bat, Kerivoula furva, forms small groups and uses furled leaves of banana (Musa formosana) as day roosts in subtropical Taiwan. In this study, we reported on the roosting ecology and social organization of K. furva. We examined whether ecological constraints, demographic traits, and physiological demands contributed to its sociality. From July 2014 to May 2016, we investigated the daily roost occupation rate, group size, and composition of each roost, and we calculated association indices in pairs. The results showed K. furva lived in groups throughout the year, and the average daily roost occupation rate was approximately 6.7% of all furled leaves that were suitable for roosting. The size of roosting groups of adults in each roost varied between 1 and 13; group size was independent of air temperature during both reproductive and nonreproductive seasons. The vast majority of roosting groups was composed of females and their young, and males frequently roosted solitarily or in a bachelor group. Forty adult bats were captured ≥4 times during the study period. The association indices in pairs of these 40 bats ranged between 0 and 0.83 with an average of 0.05 ± 0.14 (n = 780). The average association index of female–female pairs was significantly higher than that of female–male pairs and male–male pairs. Based on the association indices, the 40 bats were divided into seven social groups with social group sizes that varied between 2 and 10. Despite changing day roosts frequently, the relatively stable social bonds were maintained year‐round. Our results that groups of K. furva were formed by active aggregation of multiple generation members supported the demographic traits hypothesis.     

Male Seminole Bat Winter Roost-Site Selection in a Managed Forest

Abstract: Understanding year-round roost-site selection is essential for managing forest bat populations. From January to March, 2004 to 2006, we used radiotelemetry to investigate winter roost-site selection by Seminole bats (Lasiurus seminolus) on an intensively managed landscape with forested corridors in southeastern South Carolina, USA. We modeled roost-site selection with logistic regression and used Akaike's Information Criterion for small samples (AIC_c) and Akaike weights to select models relating roost-site selection to plot- and landscape-level variables. We tracked 20 adult male bats to 71 individual roosts. Bats used a variety of roosting structures, including the canopy of overstory trees, understory vegetation, pine (Pinus spp.) needle clusters, and leaf litter. Roost height, structure type, and habitat type were influenced by changes in minimum nightly temperature. On warmer nights, bats selected taller trees in mature forest stands, but when minimum nightly temperatures were <4° C, bats typically were found roosting on or near the forest floor in mid-rotation stands. We recommend avoiding prescribed burning in mid-rotation stands on days when the previous night's temperature is <4 °C to minimize potential disturbance and direct mortality of bats roosting on or near the forest floor. We encourage forest managers to incorporate seasonal changes in roost-site selection to create year-round management strategies for forest bats in managed landscapes.     

Comparative Roosting Ecology of Cynopterus (Chiroptera: Pteropodidae) Fruit Bats in Peninsular Malaysia1

Although the use of modified roosts has been reported in more than 20 species of bats in the tropics, comparative studies of the roosting ecology of congeneric tent‐roosting species are notably lacking. In the Paleotropics, this unique behavior has been described in two species belonging to the genus, Cynopterus: C. sphinx and C. brachyotis. However, it is not known whether tent roosting is an essential component of their roosting ecology, or whether the behavior is found in other members of the genus. In this study we characterize the roosting ecology of four sympatric species of Cynopterus in peninsular Malaysia and use these data to address two main questions. (1) Do all four species use modified roosts and, in those that do, is tent‐roosting obligate or opportunistic? (2) Do species pairs overlap in roost preferences and roosting habitat and, if so, is there evidence for interspecific interactions in relation to these resources? We radio‐tracked bats at two floristically distinct sites and located a total of 249 roosts. Interspecific roost niche overlap was minimal at both sites and we found no evidence for interspecific competition for roost resources at the local level. Species differences in roosting ecology were defined primarily by spatial separation of roosting habitats and secondarily by within‐habitat differences in roost selection. Importantly, we found that although periodic use of modified roosts was a characteristic shared by all four species, most roosts were unmodified, indicating that tent roosting is a facultative behavior in Malaysian Cynopterus.     

Communal Roost Site Selection in a Neotropical Harvestman: Habitat Limitation vs. Tradition

Many species have been reported to form roosting (resting, sleeping) aggregations at ‘traditional’ sites, but the alternative hypothesis that specific sites are used repeatedly because of habitat limitation is rarely tested. We studied the roosting behavior of a species of harvestman (Opiliones, Prionostemma sp.) at a lowland rainforest site in Nicaragua. Both sexes roosted by day in spiny palm trees, dispersed at dusk to forage, and rejoined aggregations just before dawn. The distribution of harvestmen among spiny palms was significantly clumped, and harvestman density did not correlate with spiny palm density. Aggregations formed repeatedly in a small subset of the available spiny palms and the same sites were used in two different years (2001, 2003). Nevertheless, the membership of aggregations was fluid; individual harvestmen were found at multiple roosts and moved up to 0.2 km per night. Translocated animals often returned to the roost where they had been released or nearby roosts but were never found at previously unused sites. The high consistency of site use but low site fidelity of individuals suggests that roost sites differed conspicuously (to the harvestmen) from sites that were not used. We found no univariate or multivariate differences between used and unused sites, however, in the characteristics of the trees or microclimate. These results conflict with the habitat limitation hypothesis but are consistent with the traditional site use hypothesis. The tradition may be mediated by a site‐labeling chemical, a mechanism that does not require individual site fidelity. We discuss these results in relation to the proposed functions of roosting aggregations.     

One for all and breeding for one: Cooperation and competition as a tamarin reproductive strategy

In this paper I examine behavioral strategies used by male and female tamarins to increase individual reproductive opportunities while continuing to maintain a high level of group cohesion and social cooperation. Tamarins of the genus Saguinus are characterized by social groups generally composed of more than one adult of each sex, but a breeding system in which only a single female in each group gives birth. The breeding sovereignty of a single dominant female limits the reproductive opportunities of subordinate females as well as the reproductive opportunities of resident adult males.^1-3 Despite extreme variability in year-to-year reproductive success among members of the same social group, field studies indicate that within-group intrasexual aggression and fighting are rare, and that both breeding and nonbreeding individuals expend time and energy cooperatively caring for young, defending productive feeding sites, and assisting in food harvesting activities.^4-11 In fact, Caine¹² has argued that “co-operation, tolerance, and flexibility” are the primary themes of tamarin social interactions (p. 218). A major question that remains unanswered, however, is how such high levels of cooperation could have evolved in a social system characterized by emigration of both adult males and females from the natal group, polyandrous mating, and intense reproductive competition.     

Mixed roosts in the Montagu’s harrier Circus pygargus during courtship

Mixed communal roosting of Montagu’s harrier Circus pygargus in the pre-laying period was observed on Calcareous Marshes in Eastern Poland from 1992 to 1995. To my knowledge, this behaviour was described in literature for the first time. The communal roosting in Montagu’s harrier during courtship can help in estimation of mate attraction and finally in mate choice. Harriers from communal roosts start egg laying earlier when compared to the outside roosts. Communal roosting as anti-predator behaviour can help with predator detection and provides benefits to all members of the group. The pair formation process has led to disintegration of communal roosting. Males were more common in the roosting places than females. The time of roosting was correlated with the photoperiod. The weather and predators impact delayed the formation of mixed roosting places.     

The dawn flight of the gold swift Hepialus hecta: predator avoidance and the integration of complex lek behaviour (Lepidoptera,Hepialidae)

Predator pressure is a considerable evolutionary force. The evening twilight flight of species in the family Hepialidae Swift moths has been attributed to the moths (which have no hearing and so cannot detect bats) flying in a bat/bird free window. Several species deploy elaborate lek behaviour in this period. The expected flight in the dawn twilight is little reported, except in the non‐leking Hepialus (Korscheltellus) gracilis. A detailed study of the dawn flight in the leking species Hepialus (Phymatopus) hecta shows that it is less extensive than the evening lek flight (both in duration and in the number of moths participating), is confined to a much narrower window around sunrise, involves no reproductive behaviour, and functions only to re‐locate the members of copulating pairs and sessile displaying males from the emergent ground vegetation to less conspicuous roosting sites. Compilation of individual biographies over the 24‐h cycle permits a full construction of the diel activity in time and space. It consists of two twilight flight periods, separated by prolonged roosting during the hours of darkness and daylight. The moths use almost the whole available range of roosting sites, from the base of the ground vegetation to the tree canopy, and cyclically leave and re‐enter the lek site from these positions. The copulating posture and position avoid mammalian predators, and facilitate escape from spiders and wasps by the efficient use of a dead drop. The narrowness of the dawn flight is attributed to the need for this prolonged but conspicuous copulation, which precludes a morning mating, and to a demonstrated asymmetry of twilight activity in birds. The whole integrated spatio‐temporal cycle is attributable to evolutionary pressure to minimize the impact of predators, demonstrated to include birds, bats, wasps, and spiders, and probably also dragonflies, mice and shrews; the pre‐existing dawn flight is permissive to the evolution of sessile male displays during the evening lek. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 305–319.     

Roost preferences and foraging ranges of the eastern forest bat Vespadelus pumilus under two disturbance histories in northern New South Wales,Australia

Little is known about the habitat requirements of Australian bats; however, this information is needed to make better‐informed decisions when systems are disturbed. This study contrasts the roosting and foraging ecology of the eastern forest bat Vespadelus pumilus (Vespertilionidae), one of Australia’s smallest bats, between two sites of differing disturbance history on the mid‐north coast of New South Wales. Lorne Flora Reserve (182 ha) is primarily old‐growth forest surrounded by regrowth forest and eucalypt plantations, while Swans Crossing is dominated by regrowth and eucalypt plantations established on part of an old dairy farm. A total of 38 bats were tracked during the maternity and mating seasons at the two sites. Roost preferences were determined by comparing trees used as roosts with those randomly available, while foraging bats were triangulated from fixed stations at night. Bats tracked at Lorne Flora Reserve typically roosted in hollows within large, mature trees and showed a strong preference for roosting and foraging (females only) within the Reserve. Lactating females at Swans Crossing roosted in hollows of remnant rainforest trees within a gully and dead eucalypts, while males often roosted in understorey trees (such as Acacia). Dead trees were frequently used as roosts at both sites. Under both disturbance histories, the mean distance of female maternity roosts from creeks was 20 m, indicating that riparian zones provide important roosting habitat for V. pumilus. However, roosts shifted to the mid‐slope prior to winter when bats mate. Retention of mature trees in a variety of topographic locations may allow behavioural adjustments with the seasons. Bats caught in the regrowth forest also foraged there, with foraging ranges averaging just 5.3 ha (n = 10), indicating that regrowth is used by this bat for both foraging and roosting.     

Bats are able to maintain long-term social relationships despite the high fission-fusion dynamics of their groups

Elephants, dolphins, as well as some carnivores and primates maintain social links despite their frequent splitting and merging in groups of variable composition, a phenomenon known as fission-fusion. Information on the dynamics of social links and interactions among individuals is of high importance to the understanding of the evolution of animal sociality, including that of humans. However, detailed long-term data on such dynamics in wild mammals with fully known demography and kin structures are scarce. Applying a weighted network analysis on 20,500 individual roosting observations over 5 years, we show that in two wild Bechstein's bat colonies with high fission-fusion dynamics, individuals of different age, size, reproductive status and relatedness maintain long-term social relationships. In the larger colony, we detected two stable subunits, each comprising bats from several family lineages. Links between these subunits were mainly maintained by older bats and persisted over all years. Moreover, we show that the full details of the social structure become apparent only when large datasets are used. The stable multi-level social structures in Bechstein's bat colonies resemble that of elephants, dolphins and some primates. Our findings thus may shed new light on the link between social complexity and social cognition in mammals.