Tent Construction by the Short-nosed Fruit Bat Cynopterus sphinx (Chiroptera: Pteropodidae) in Southern India

The short-nosed fruit bat Cynopterus sphinx (Chiroptera: Pteropodidae) constructs shelters by severing stems of the curtain creeper, Vernonia scandens, and stems and leaves of the mast tree, Polyalthia longifolia, creating partially enclosed cavities (stem tents) in which to roost. Our observations indicate that the construction and maintenance of stem tents are primarily, if not exclusively, the behaviour of single males. A stem tent is formed in V. scandens when a single male C. sphinx severs up to 300 small- to medium-sized stems creating a partially flattened, bell-shaped cavity, and in P. longifolia when a male severs a few medium- to small-sized branches and many leaf petioles, creating an entry/exit portal and space in which to roost. A tent constructed in V. scandens is completed in approximately 30 d, whereas one in P. longifolia is completed in about 50 d. Stem-tent construction takes place mostly at night, but some stem chewing occurs in late afternoon. At night a stem tent is occupied by a single male, whereas females are usually absent. During the day the number of bats occupying completed tents is highly variable, ranging from two to 19 females (and their pups) and a single adult male. Tent construction is annually bimodal, which corresponds to a biannual breeding season. A dominant male sometimes deposits saliva on branches inside his tent cavity and actively defends this space from intrusions by other males. Both behaviours suggest forms of scent marking and territorial display. Our observations indicate that dominant males construct tents, recruit females and then defend the tents (and their female occupants) for the purpose of gaining reproductive access. The variance in harem group size indicates that some tent-making males are more successful than others in recruiting females.     

犬蝠夜栖息地及夜栖息巢特征的初步研究

<正>大多数种类的蝙蝠不会整个晚上都进行觅食,通常在觅食期间有一段长短不一的时间停留在临时地休息,此为夜栖息行为(Hatfield,1937;Krutzsch,1954;Barbour and Davis,1969;Kunz,1973,1974;Hirshfeld et al.,1977)。蝙蝠在夜栖息地进食(Vaughan,1976;Funakoshi and Maeda,2003)、休息并消化食物(Brigham,1991;Funakoshi and Maeda,2003),甚至社会交流(Kunz,1982;Kunz and Lumsden,2003)。不同种类的蝙蝠     

Convergence in tent architecture and tent-making behavior among neotropical and paleotropical bats

Fifteen species of neotropical and three species of paleotropical bats are known either to roost in or to make tents in over 80 species of vascular plants. We summarize the current knowledge of bat-tent architecture, report two new styles of tents (conical and apical) from the Paleotropics, compare the similarity in tents constructed, or used, by neotropical and paleotropical bats, and consider possible functions of tents. Seven styles of tents are known from the Neotropics, three (conical, palmate umbrella, and apical tents) are known from both the Neo- and the Paleotropics, and one (stem tent) is unique to the Paleotropics. In the Neotropics tent-roosting and/or tent-making appears to be a behavior unique to the diverse microchiropteran family Phyllostomidae (subfamily Phyllostomatinae: tribe Stenodermatini), and in the Paleotropics two members of the megachiropteran family Pteropodidae and one member of the microchiropteran family Vespertilionidae are known to construct or roost in tents. Despite the variety of plant taxa used by bats in tent construction, there appears to be a limited number of different leaf forms that can be altered by bats and used as tents. We suggest that the similarity in tent architecture observed among the neotropical and paleotropical bats is a consequence of convergence in leaf morphology among forest understory plants. The congruence in tent-making/roosting behavior observed in members of the Stenodermatini and the Pteropodidae (genusCynopterus) suggests a phylogenetic influence on these behaviors. The similarity in tent-making and/or tent-roosting behavior and life-history traits (small, <70 g, mostly foliage-roosting frugivores) among these divergent neotropical and paleotropical taxa supports a convergence hypothesis in which members of these groups have become ecological equivalents. Although actual tent-making has been observed in only one bat species to date, we suggest that the principal selective force leading to the evolution of tent-making is a polygynous mating system whereby males construct tents to gain access to females. Tents in turn provide resources that offer protection from predators and inclement weather.     

Roosting patterns in a captive colony of short-nosed fruit bat Cynopterus sphinx (Vahl)

Development of roosting patterns under a limited resource was studied in the short-nosed fruit bat C. sphinx in captivity. Spatial fidelity during the resting period (day time) and the individual male bat's presence/absence in the roost (occupancy index) were estimated during the active period (night time). Results show the presence of three groups on the basis of spatial fidelity. The first group was associated with the tent consisting of a harem male and seven females. The second group stayed near to the harem. The third group consisting of two males showed little occupancy index and no spatial fidelity. Female turnover between the first and second groups, and harem male replacement were observed. These findings of male groupings and female loyalty on the basis of "resource", suggest that resource defence polygyny is the primary mating strategy in C. sphinx.     

海口地区犬蝠冬季食性及栖宿地类型

犬蝠(Cynopterus sphinx)是海口地区较为常见的一种果蝠,属于近危兽类。对其生态学研究有助于了解犬蝠在生态系统中的作用,并为野生动物管理提供基础资料。从2005年10月至2006年1月,采用直接观察法、拾遗法、粪便分析法并结合雾网采样法对本地区犬蝠冬季的栖宿地类型及食性进行了研究。结果发现,犬蝠主要栖宿在椰子等棕榈科植物的大型叶片下以及废弃的房屋中;其食物包括9科11种植物的果实、2科2种植物的叶片和1种植物的茎。     

Effect of female group size on harem male roosting behavior of the Indian short-nosed fruit bat <Emphasis Type="Italic">Cynopterus sphinx</Emphasis>

Mate guarding has been known to incur costs and cause constraints for harem males in many polygynous species. However, the effect of female group size on the harem male’s time budget in bats has received very limited attention. The Indian short-nosed fruit bat, Cynopterus sphinx, exhibits resource defense polygyny, in which tent roosting males construct tents and defend multiple female bats. We studied the effect of female group size on three aspects of harem male behavior: social grooming by reciprocal licking, tent maintenance, and tent guarding in the mast tree Polyalthia longifolia. In the process of reciprocal licking, all the bats in the harem were drenched in saliva before emergence, and this activity was positively and significantly correlated with female group size. Once females departed for foraging, harem males remained in their respective tents at night-time between intermittent foraging bouts and engaged in tent maintenance and tent guarding. Time invested by harem male bats in tent maintenance and tent guarding were positively and significantly correlated with female group size. Harem males extended their presence in tent by utilizing tents as feeding roosts. Female group size also influenced the emergence time of harem male bats, where males with largest group emerged later than did the smallest group. Likewise, harem male with the smallest group had more time available for foraging than the male with the largest group. Findings of this study suggest that having a larger harem may indeed be costly for the males by reducing their foraging time.     

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.     

Roosting ecology of Stenodermatinae bats (Phyllostomidae): evolution of foliage roosting and correlated phenotypes

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Contrasting genetic structure in two co-distributed species of old world fruit bat

The fulvous fruit bat (Rousettus leschenaulti) and the greater short-nosed fruit bat (Cynopterus sphinx) are two abundant and widely co-distributed Old World fruit bats in Southeast and East Asia. The former species forms large colonies in caves while the latter roots in small groups in trees. To test whether these differences in social organization and roosting ecology are associated with contrasting patterns of gene flow, we used mtDNA and nuclear loci to characterize population genetic subdivision and phylogeographic histories in both species sampled from China, Vietnam and India. Our analyses from R. leschenaulti using both types of marker revealed little evidence of genetic structure across the study region. On the other hand, C. sphinx showed significant genetic mtDNA differentiation between the samples from India compared with China and Vietnam, as well as greater structuring of microsatellite genotypes within China. Demographic analyses indicated signatures of past rapid population expansion in both taxa, with more recent demographic growth in C. sphinx. Therefore, the relative genetic homogeneity in R. leschenaulti is unlikely to reflect past events. Instead we suggest that the absence of substructure in R. leschenaulti is a consequence of higher levels of gene flow among colonies, and that greater vagility in this species is an adaptation associated with cave roosting.     

The dynamics of tent‐roosts in the palm Sabal mauritiiformis and their use by bats in a montane dry forest

Tent‐making bats modify leaves to build refuges. Leaf modification involves energetic and defense costs that should be balanced by the benefits of tent‐roosting. The alteration of the leaf's vascular system reduces the tent's life expectancy, so to obtain a benefit, bats are expected to use tents regularly as long as they remain functional and not modify more leaves than necessary. Over 2 yr, we documented the dynamics of tent construction and use by Uroderma convexum and other bat species in the palm Sabal mauritiiformis in a Colombian transitional dry forest. We also assessed tent condition and compared it to nonmodified leaves of approximately the same age in focal palms. Probability of tent use by U. convexum varied between 57 percent during a reproductive period and 4 percent outside of this period. Bats cut the main vein of folioles, partially affecting water transport in the leaf. However, there were no differences between tents and nonmodified leaves in deterioration scores or deterioration rates over 1 yr. During 2 yr, 48 tents were lost for different causes, but this loss was balanced by the construction of 51 new tents. Thus, bats maintained an excess of usable tents. Palm leaves are long‐lived and seem preadapted to sustain damage and remain viable, particularly in species growing in dry environments. We present several hypotheses to explain the advantage of maintaining a tent surplus.     

First application of satellite telemetry to track African straw-coloured fruit bat migration

Despite long-standing awareness of the potentially important ecological role of fruit bats, we know little about the ecology of the vast majority of species. Here we report the results of a pilot satellite tracking study aimed at establishing the scale of movement of the straw-coloured fruit bat Eidolon helvum . This was the first ever attempt to track African fruit bats using satellite telemetry. We tagged four bats with solar-charged 12 g satellite transmitters at Kasanka National Park in December 2005 and obtained a combined total of 104 different location fixes over a 149-day period. Before migrating, bats foraged as far as 59 km from the roost in a single evening; by contrast, one migrating individual moved 370 km in one night. Bats travelled an average 29 km day⁻¹ over the period of study, with bats that appeared to be migrating moving north-west from Kasanka at an average 90 km day⁻¹. The greatest cumulative distance travelled by a single bat was 2518 km in 149 days. The results show conclusively that the straw-coloured fruit bat E. helvum is capable of migrating thousands of kilometres across central Africa on an annual basis, implying that the fruit pulse in northern Zambia is richer than anything on offer in the Democratic Republic of the Congo at the same time of the year.     

The effect of logging on fission-fusion behaviour of tree-dwelling bats explored by an agent-based model

Logging is one of the greatest threats to global biodiversity, while forests are one of the most important habitats for bats. Bats that roost in tree cavities require a large number of potential roosts due to their frequent roost switching. However, the density of tree cavities and hollows sufficient to sustain large populations of bat species in forests is unknown. The fission-fusion dynamics of bat groups in forest environment is associated with ritualised dawn swarming behaviour at potential tree cavities that serves to exchange information in a non-centralised decision-making process. We used a computer model based on the swarm algorithm, SkyBat, that resembles this complex process and aimed to determine how population size changes over time when cavity trees are removed from roosting territory of the local population of Leisler's bats (Nyctalus leisleri), which inhabit a forest habitat in Central Europe. Simulations revealed that social bonds between bats, maintained by frequent switching among groups, play an important role in this highly dynamic system. When strong social contact was not considered, reducing the original number of trees with cavities (20 cavities × ha⁻¹) to 50% was still acceptable to bats, but further interventions and/or increased demand for social contact would have led to local extinction of the species. Results suggest that potential bat roosts in mature forest stands should be preserved as much as possible and that non-intensive logging and management can be beneficial to tree-dwelling bats.     

Seed Dispersal of Morus macroura (Moraceae) by Two Frugivorous Bats in Xishuangbanna, SW China

The dispersal of Morus macroura seeds by two species of frugivorous bats ( Rousettus leschenaulti and Cynopterus sphinx ) was studied in a forest at Xishuangbanna Tropical Botanical Garden in Southwest China from March to May 2005. Feeding roosts were identified within 500 m around parent trees and the types and number of seed loads under each roost were recorded. We found feeding roost density decreased with increasing distance from the parent, but found no correlation between distance and seed deposition. The effect of bat digestion on seed germination was investigated, and we found that germination percentage of all treatments involving ingestion by bats was significantly lower than control seeds and some germination parameters of seeds from different treatments changed.     

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.     

Tent selection, roosting ecology and social organization of the tent-making bat, Ectophylla alba, in Costa Rica

Small groups of the tent-making bat, Ectophylla alba , were found roosting in Heliconia (Musaceae) tents in old secondary-growth forest in north-eastern Costa Rica. The choice of specific Heliconia leaves for tents was predicted on the basis of leaf size and age. Additionally, tents in shrubs, saplings and epiphytic plants were found scattered throughout both primary- and secondary-growth forest. Tents were used either as night feeding roosts or as day-roosts for as long as 45 days. Groups of bats remained together when they moved to newly cut tents. After parturition, tent groups divided into all-male colonies and maternity colonies with females, non-volant young and a single adult male.     

Calls in the Forest: A Comparative Approach to How Bats Find Tree Cavities

Although tree cavities are a particularly critical resource for forest bats, how bats search for and find new roosts is still poorly known. Building on a recent study on the sensory basis of roost finding in the noctule (Ruczynski et al. 2007), here we take a comparative approach to how bats find roosts. We tested the hypothesis that species' flight abilities and echolocation call characteristics play important roles in how well and by which cues bats find new tree roosts. We used the very manoeuvrable, faintly echolocating brown long-eared bat ( Plecotus auritus ) and the less manoeuvrable, louder Daubenton's bat ( Myotis daubentonii ) as study species. The species are sympatric in European temperate forests and both roost in tree cavities. We trained bats in short-term captivity to find entrances to tree cavities and experimentally manipulated the sensory cues available to them. In both species, cue type influenced the search time for successful cavity detection. Visual, olfactory and temperature cues did not improve the bats' performance over the performance by echolocation alone. Eavesdropping on conspecific echolocation calls played back from inside the cavity decreased search time in Daubenton's bat ( M. daubentonii ), underlining the double function of echolocation signals – orientation and communication. This was not so in the brown long-eared bat ( P. auritus ) that has low call amplitudes. The highly manoeuvrable P. auritus found cavities typically from flight and the less manoeuvrable M. daubentonii found more entrances during crawling. Comparison with the noctule data from Ruczyński et al. (2007) indicates that manoeuvrability predicts the mode of cavity search. It further highlights the importance of call amplitude for eavesdropping and cavity detection in bats.     

犬蝠对不同浓度香蕉汁的选择

为了解犬蝠的嗅觉灵敏度,我们以6 只成年犬蝠对不同浓度的成熟香蕉汁和纯水对照的选择倾向和取食时间各进行了连续5 个夜晚的室内观察。香蕉汁的浓度分别为100% (120 g),50% (60 g 香蕉汁和60 g 纯水),25% (30 g 香蕉汁和90 g 纯水),12. 5% (15 g 香蕉汁和105 g 纯水);另取纯水(W)与熟烂的香蕉汁(OR)各120 g 作为对照。将以上食物和对照随机注入不同的容器内,观察犬蝠对不同浓度的香蕉汁、熟烂香蕉汁和纯水的选择。结果表明,犬蝠对纯成熟香蕉汁的选择性最强,取食次数和取食量均最高,而停留时间也最长;说明犬蝠具有分辨不同浓度的气味差异。我们推测犬蝠在野外觅食时可能优先取食成熟水果。     

Roosting Requirements of Two Frugivorous Bats (Sturnira lilium and Arbiteus intermedius) in Fragmented Neotropical Forest1

As tropical forest fragmentation accelerates, scientists are concerned with the loss of species, particularly those that play important ecological roles. Because bats play a vital role as the primary seed dispersers in cleared areas, maintaining healthy bat populations is critical to natural forest regeneration. Observations of foraging bats suggest that many Neotropical fruit‐eating species have fairly general habitat requirements and can forage in many different kinds of disturbed vegetation; however, their roosting requirements may be quite different. To test whether or not general foraging requirements are matched by equally broad roosting requirements, we used radiotelemetry to locate roost sites of two common frugivorous bat species (Sturnira lilium and Artibeus intermedius) in a fragmented forest in southeastern Mexico. Sturnira lilium roosted inside tree cavities and selected large‐diameter roost trees in remnant patches of mature forest. Fewer than 2 percent of trees surveyed had a mean diameter equal to or greater than roost trees used by . S. lilium, Artibeus intermedius roosted externally on branches and vines and under palm leaves and selected roost trees of much smaller diameter. Compared to random trees, roost trees chosen by A. intermedius were closer to neighboring taller trees and also closer in height to these trees. Such trees likely provide cryptic roosts beneath multiple overlapping crowns, with sufficient shelter from predators and the elements. While males of A. intermedius generally roosted alone in small trees within secondary forest, females roosted in small groups in larger trees within mature forest and commuted more than three times farther than males to reach their roost sites. Loss of mature forest could impair the ability of frugivorous bats to locate suitable roost sites. This could have a negative impact on bat populations, which in turn could decrease forest regeneration in impacted areas.     

Predation by squirrel monkeys and double-toothed kites on tent-making bats

Central American squirrel monkeys (Saimiri oerstedi) appear to recognize the modified leaves that phyllostomid bats utilize for diurnal roost sites. The monkeys visually and manually search these bat tents for both bats and insects. Adult males are the most successful at capturing bats. Nonvolant juvenile bats are more vulnerable to monkey predation than are adults. Bats that escape monkey predation frequently are captured by doubletoothed kites (Harpagus bidentatus) that tend foraging troops of monkeys. Predation by squirrel monkeys, coupled with that of double-toothed kites, may be a significant source of mortality for tent-making bats.     

广州市犬蝠对栖息地的选择

2006年9月-2007年3月,在广州市区对犬蝠(Cynopterus sphinx angulatus)的分布和栖息地的选择进行了研究.结果表明:共发现犬蝠栖巢45个,有犬蝠栖息的26个;犬蝠对栖息植物的选择具有专一性,主要利用景观树蒲葵(Livistona chinensis)的叶片建造栖巢,其栖息地跟人类活动区域高度重合.捕获的18个栖群中5个栖群为全捕,13个栖群有部分个体未捕获;其中17个栖群中包括有雄性个体,占群体总数的94%;5个全捕栖群中,2个为一雄多雌,一个为一雄一雌群体,一个为独栖雄性,另一个为多个雄性的"单身汉群体".