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.     

Day roost selection in female Bechstein's bats (Myotis bechsteinii): a field experiment to determine the influence of roost temperature

The decision where to live has far-reaching fitness consequences for animals. In contrast to most other mammals or birds that use sheltered nest sites, female Bechstein's bats frequently switch day roosts during one breeding season, and therefore must often decide where to spend the day. Selecting the right roost is important, because roost quality, e.g. microclimatic condition, influences survival and reproduction in bats. Although thermal factors are very important for the quality of roosts occupied by bats, whether bats base their day roost selection directly on roost temperature has not been tested in the field. Over one summer, we examined and tested the roost choice of 21 individually marked female Myotis bechsteinii living in one maternity colony. In a field experiment, we allowed the bats to choose between relatively warm versus cold bat boxes, while controlling for site preferences. We expected females to exhibit a preference for warm roosts during pregnancy and lactation to accelerate gestation and shorten the period of growth of their young. Roost occupancy over 160 census days reflected significant temperature differences among 89 surveyed roosts (14 tree holes and 75 bat boxes), and preferences changed with the season. Females significantly preferred cold roosts before parturition, whereas post-partum, they significantly favoured warm roosts. Temperature preferences were independent of the roost site, and thus roost selection was based directly on temperature. Boxes with significantly different daytime temperatures did not differ significantly at night. Consequently, bats would have to spend at least 1 day in a new roost to test it. Information transfer among colony members might facilitate knowledge of roost availability. Access to many roosts providing different microclimates is likely to be important for successful reproduction in the endangered Bechstein's bat.     

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.     

Migratory behaviour of the Schreiber's bat: when, where and why do cave bats migrate in a Mediterranean region?

Regional migrations are important elements of the biology of bats, but remain poorly understood. We obtained a large dataset of recoveries of ringed Miniopterus schreibersii to study the patterns and drivers of migration of a Mediterranean cave-dwelling bat. In spite of the mildness of Mediterranean winters, in average years bats hibernated, and few movements were recorded during this period. After hibernation, females migrated to spring roosts, and again to maternity roosts just before parturition. This late arrival at nurseries could be a strategy to avoid a harmful build-up of parasites. Soon after the juveniles were weaned, the mothers migrated to the roosts where they spent autumn and sometimes also winter. Juveniles remained in the warm nurseries longer, presumably because high roost temperatures speed up growth. The pattern of migration of males was similar to that of females, but they left hibernacula later and remained more mobile during the maternity season. They also arrived at the hibernacula later, possibly because they needed time to build up fat stores after the energetically costly mating season. Maternity colonies spent the yearly cycle in well-defined home ranges (mean=19 030 km²), which overlapped greatly. Bats were furthest from the maternity sites during hibernation, but even then 80% remained within 90 km of them. Each hibernaculum attracted bats from multiple nurseries, from within a mean range of 10 770 km². We tested two potential drivers for migration – temperature in the roosts and at the foraging areas – but our results supported only the first one. Bats migrated to reach the roosts most thermally suited for each phase of their life cycle, indicating that roost temperature and associated metabolic advantages are key drivers for regional migrations of cave-dwelling bats.     

Information transfer about roosts in female Bechstein's bats: an experimental field study

Information transfer among group members is believed to play an important part in the evolution of coloniality in both birds and bats. Although information transfer has received much scientific interest, field studies using experiments to test the underlying hypotheses are rare. We used a field experiment to test if communally breeding female Bechstein's bats (Myotis bechsteinii) exchange information regarding novel roosts. We supplied a wild colony, comprising 17 adult females of known relatedness, with pairs of suitable and unsuitable roosts and monitored the arrival of individuals marked with transponders (PIT-tags) over 2 years. As expected with information transfer, significantly more naive females were recruited towards suitable than towards unsuitable roosts. We conclude that information transfer about roosts has two functions: (i) it generates communal knowledge of a large set of roosts; and (ii) it aids avoidance of colony fission during roost switching. Both functions seem important in Bechstein's bats, in which colonies depend on many day roosts and where colony members live together for many years.     

Effects of Hierarchical Roost Removal on Northern Long-Eared Bat (Myotis septentrionalis) Maternity Colonies

Forest roosting bats use a variety of ephemeral roosts such as snags and declining live trees. Although conservation of summer maternity habitat is considered critical for forest-roosting bats, bat response to roost loss still is poorly understood. To address this, we monitored 3 northern long-eared bat (Myotis septentrionalis) maternity colonies on Fort Knox Military Reservation, Kentucky, USA, before and after targeted roost removal during the dormant season when bats were hibernating in caves. We used 2 treatments: removal of a single highly used (primary) roost and removal of 24% of less used (secondary) roosts, and an un-manipulated control. Neither treatment altered the number of roosts used by individual bats, but secondary roost removal doubled the distances moved between sequentially used roosts. However, overall space use by and location of colonies was similar pre- and post-treatment. Patterns of roost use before and after removal treatments also were similar but bats maintained closer social connections after our treatments. Roost height, diameter at breast height, percent canopy openness, and roost species composition were similar pre- and post-treatment. We detected differences in the distribution of roosts among decay stages and crown classes pre- and post-roost removal, but this may have been a result of temperature differences between treatment years. Our results suggest that loss of a primary roost or ≤ 20% of secondary roosts in the dormant season may not cause northern long-eared bats to abandon roosting areas or substantially alter some roosting behaviors in the following active season when tree-roosts are used. Critically, tolerance limits to roost loss may be dependent upon local forest conditions, and continued research on this topic will be necessary for conservation of the northern long-eared bat across its range.     

The potential availability of roosting sites for lesser short‐tailed bats (Mystacina tuberculata) on Kapiti Island,New Zealand: Implications for a translocation

Abstract

Lesser short‐tailed bats (Mystacina tuberculata) have recently been translocated to Kapiti Island in an attempt to form a new population of this threatened species. However, the island's vegetation is regenerating, and there was doubt that the forests provided enough large trees with cavities for bats to roost in. This study measured the availability of tree‐trunk cavities of the right size for potential roost sites on Kapiti Island, and assessed if habitat restoration would be required to increase the translocation's chance of success. first, trees with cavities accessible to us were sampled in six of Kapiti Island's forest types. Size variables known to affect roost site selection by lesser short‐tailed bats at the tree and cavity level were measured. Trees were classified as containing cavities that could potentially provide suitable roosts if their values for all variables measured fell within the range of roosts used by lesser short‐tailed bats in natural populations. Roosts were classified as suitably sized for solitary bats or for colonies, using measurements from both types of roosts in natural populations. Second, the density of these potential roost cavities was calculated. Cavities of a size potentially suitable for colonies were found in four of the six forest types at densities ranging from 3.2 ± 3.2 Se to 52.4 ± 14.0 trees per ha. density of potential solitary roosts was much higher. Not all potential cavities will be suitable because they may be damp, poorly insulated, or have an unsuitable microclimate. Nevertheless, our estimates indicated that the two most extensive forest types each contained thousands of potential cavities of a size suitable for colonies of lesser short‐tailed bats. In addition, there were tens of thousands of cavities large enough to shelter solitary bats. Roost habitat restoration appears unnecessary to assist translocated Mystacina tuberculata on Kapiti Island.     

Population structure of Daubenton’s bats is responding to microclimate of anthropogenic roosts

Roost microclimate plays an important role in the survival, growth and reproduction in microbats. Entering torpor is one of the main energy saving mechanisms commonly used by microbats. The use of torpor is affected by roost microclimate and seasonally differs between the two sexes in relation to their reproductive condition. Consequently, thermal properties of male and female roosts should differ. To test this hypothesis, we compared temperature parameters of two anthropogenic day roosts of Daubenton’s bats with a different structure of the population inhabiting them. In accordance with our predictions, the roost occupied by a male-dominated colony was colder and more fluctuant than the maternity roost with a female-dominated population. However, using of the two roosts changed during the season in response to changing energetic demands of the two sexes. While males were almost absent in the warmer maternity roost during pregnancy and lactation, they appeared in this roost during the post-lactation when mating starts. In contrast, females did not use the colder (male) roost until the time of weaning of juveniles, i.e., the time when their thermoregulatory needs change and they may benefit from using colder roost. Our study provides the evidence that the same roost may be used by individuals of different sex and reproductive state in different periods of the year. Generalizations about roost selection without knowledge of temporal variation in roost use and microclimatic conditions should be taken with caution. Anthropogenic roosts may be advantageous to Daubenton’s bats as these can provide a variety of suitable microclimates and/or more space for roosting than tree cavities.     

Summer tree roost selection by Rafinesque's big-eared bat

Roost requirements of most North American forest bats are well-documented, but questions remain regarding the ultimate mechanisms underlying roost selection. Hypotheses regarding roost selection include provision of a stable microclimate, space for large colonies, protection from predators, and proximity to foraging habitat, among others. Although several hypotheses have been proposed, specific mechanisms likely vary by species and geographic region. Rafinesque's big-eared bat (Corynorhinus rafinesquii) commonly roosts in trees with large basal hollows in the Coastal Plain of the southeastern United States. Our objective was to weigh evidence for hypotheses regarding selection of diurnal summer roosts by Rafinesque's big-eared bat at 8 study sites across the Coastal Plain of Georgia, USA. We used transect searches and radiotelemetry to locate roosts and measured 22 characteristics of trees, tree cavities, and surrounding vegetation at all occupied roosts and for randomly selected unoccupied trees. We evaluated 10 hypotheses using single-season occupancy models and used Akaike's information criterion to select the most parsimonious models. We located 170 tree roosts containing approximately 870 bats for our analysis. The best supported model predicted bat presence from cavity size, interior wall texture, and number of entrances. Because large cavities allow bats to fly and smooth walls impede attacks by terrestrial predators, our results are consistent with the hypothesis that bats select roosts that allow them to evade predators. However, data on predation rates are needed for a conclusive determination. Because trees suitable as roosts for Rafinesque's big-eared bat are rare in the landscape, protection of suitable forested wetland habitat is essential to provide current and long-term roost tree availability. © 2012 The Wildlife Society.     

Roost selection by Formosan leaf-nosed bats (Hipposideros armiger terasensis)

Patterns of roost use by Formosan leaf-nosed bats (Hipposideros armiger terasensis) were studied from November 1998 to April 2000. Structural characteristics, microclimates, and disturbance levels of 17 roosts used by H. a. terasensis and 15 roosts either used by other bat species (2) or not occupied by any bat species were compared. Roosts used by these bats were significantly larger in size and had greater areas covered by water compared to unused roosts. Entrances of active roosts were more likely to be east-west oriented. Hibernacula had lower entrances and ceilings than did roosts used only in summer. Higher temperatures were recorded in non-breeding roosts than in breeding roosts, but temperature gradients in these two types of roosts did not differ. In winter, hibernacula were warmer, and the temperature fluctuated less than in non-hibernacula. The relative humidities in summer roosts and hibernacula were nearly 100%. Disturbance levels were significantly higher in non-breeding roosts than in breeding roosts, and in non-hibernacula than in hibernacula. These results suggest that the Formosan leaf-nosed bats are selective of their roosts, but the pattern of their roost selection differs from those reported for bats of temperate regions. The reasons for such differences may be related to differences in body size, behavior, and reproductive strategy of the Formosan leaf-nosed bats living in a subtropical climate in Taiwan.     

Modelling Sensory Limitation: The Role of Tree Selection,Memory and Information Transfer in Bats’ Roost Searching Strategies

Sensory limitation plays an important role in the evolution of animal behaviour. Animals have to find objects of interest (e.g. food, shelters, predators). When sensory abilities are strongly limited, animals adjust their behaviour to maximize chances for success. Bats are nocturnal, live in complex environments, are capable of flight and must confront numerous perceptual challenges (e.g. limited sensory range, interfering clutter echoes). This makes them an excellent model for studying the role of compensating behaviours to decrease costs of finding resources. Cavity roosting bats are especially interesting because the availability of tree cavities is often limited, and their quality is vital for bats during the breeding season. From a bat’s sensory point of view, cavities are difficult to detect and finding them requires time and energy. However, tree cavities are also long lasting, allowing information transfer among conspecifics. Here, we use a simple simulation model to explore the benefits of tree selection, memory and eavesdropping (compensation behaviours) to searches for tree cavities by bats with short and long perception range. Our model suggests that memory and correct discrimination of tree suitability are the basic strategies decreasing the cost of roost finding, whereas perceptual range plays a minor role in this process. Additionally, eavesdropping constitutes a buffer that reduces the costs of finding new resources (such as roosts), especially when they occur in low density. We conclude that natural selection may promote different strategies of roost finding in relation to habitat conditions and cognitive skills of animals.     

Group decision making in fission-fusion societies: evidence from two-field experiments in Bechstein's bats

Group decisions are required when group coordination is beneficial, but individuals can choose between alternatives. Despite the increased interest in animal group decision making, there is a lack of experimental field studies that investigate how animals with conflicting information make group decisions. In particular, no field studies have considered the influence of fission-fusion behaviour (temporary splitting into subgroups) on group decisions. We studied group decision making in two wild Bechstein's bat colonies, which are fission-fusion societies of stable individual composition. Since they frequently switch communal roosts, colony members must regularly make group decisions over where to roost. In the two-field experiments, we provided marked individuals with conflicting information about the suitability of potential roosts. We investigated whether conflicting information led to group decisions that followed a 'unanimous' or a 'majority' rule, or increased colony fission. Individual behaviour suggests that bats considered both their own information and the behaviour of others when deciding where to roost. Group decisions about communal roosts reflected the information available to a majority of the bats roosting together, but conflicting information led to an increased fission in one colony. Our results suggest that fission-fusion societies allow individuals to avoid majority decisions that are not in their favour.     

Roosts of Allen's Lappet-Browed Bat in Northern Arizona

ABSTRACT In Arizona, USA, Allen's lappet-browed bat (Idionycteris phyllotis) forms maternity colonies in ponderosa pine (Pinus ponderosa) snags. There is little information on the roosting habitat of males. We used radiotelemetry to locate 16 maternity, 3 postlactating, and 2 bachelor roosts and combined data with unpublished data for maternity roosts (n = 11) located in 1993–1995. Most (96%) maternity roosts were in large-diameter (x̄ ± SE: 64 ± 2.7 cm) ponderosa pine snags under sloughing bark. Models that best predicted the probability of a snag's use as a maternity roost indicated bats selected taller snags closer to forest roads than comparison snags. Maternity roosts averaged 11 bats per roost (SE = 2, n = 15; from exit counts) and were an average distance of 1.6 km from capture sites (SE = 0.3, n = 17). Bachelor roosts were in vertical sandstone cliff faces in pinyon-juniper (Pinus edulis-Juniperus spp.) woodlands approximately 12 km from capture sites; these and other capture records in Arizona indicated sexual segregation may have occurred during the maternity season. Of 11 maternity snag roosts located in 1993–1995, only one continued to function as a roost. Resource managers should maintain patches of large-diameter ponderosa pine snags with peeling bark to provide maternity roosting habitat for Allen's lappet-browed bat.     

Roost use by long-tailed bats in South Canterbury: examining predictions of roost-site selection in a highly fragmented landscape

We studied the roosting ecology of the long-tailed bat (Chalinolobus tuberculatus) during the springautumn months from 1998–2002 at Hanging Rock in the highly fragmented landscape of South Canterbury, South Island, New Zealand. We compared the structural characteristics and microclimates of roost sites used by communally and solitary roosting bats with those of randomly available sites, and roosts of C. tuberculatus occupying unmodified Nothofagus forest in the Eglinton Valley, Fiordland. Roosting group sizes and roost residency times were also compared. We followed forty radio-tagged bats to 94 roosts (20% in limestone crevices, 80% in trees) at Hanging Rock. Roosts were occupied for an average of 1 day and 86% were only used once during the study period. Colony size averaged 9.8 ± 1.1 bats (range 2–38) and colonies were dominated by breeding females and young. Indigenous forest, shrubland remnants and riparian zones were preferred roosting habitats. Communally roosting bats selected roosts in split trunks of some of the largest trees available. Selection of the largest available trees as roost sites is similar to behaviour of bat species occupying unmodified forested habitats. Temperatures inside 12 maternity roosts measured during the lactation period were variable. Five roosts were well insulated from ambient conditions and internal temperatures were stable, whereas the temperatures inside seven roosts fluctuated in parallel with ambient temperature. Tree cavities used by bats at Hanging Rock were significantly nearer ground level, had larger entrance dimensions, were less well insulated, and were occupied by fewer bats than roosts in the Eglinton Valley. These characteristics appear to expose their occupants to unstable microclimates and to a higher risk of threats such as predation. We suggest that roosts at Hanging Rock are of a lower quality than those in the Eglinton Valley, and that roost quality may be one of the contributory factors in the differential reproductive fitness observed in the two bat populations. The value of introduced willows (especially Salix fragilis) as bat roosts should be acknowledged. We recommend six conservation measures to mitigate negative effects of deterioration of roosting habitat: protection and enhancement of the quality of existing roosts, replanting within roosting habitat, provision of high quality artificial roosts, predator control, and education of landowners and statutory bodies.     

Bats relocate maternity colony after the natural loss of roost trees

Understanding the ephemerality of trees used as roosts by wildlife, and the number of roost trees needed to sustain their populations, is important for forest management and wildlife conservation. Several studies indicate that roosts are limiting to bats, but few studies have monitored longevity of roost trees used by bats over several years. From 2004–2007 in Cypress Hills Interprovincial Park, Saskatchewan, Canada, several big brown bats (Eptesicus fuscus) from a maternity group roosted in cavities in trembling aspen (Populus tremuloides) trees approximately 7 km southeast away from their original known roosting area (RA1). Using a long-term data set of the roost trees used by bats in this area from 2000–2007, we evaluated whether the movement of bats to the new roosting area (RA4) corresponded with annual and cumulative losses of roost trees. We also determined whether longevity of the roosts from the time we discovered bats first using them differed between the 2 roosting areas based on Kaplan-Meier estimates. Bats began using RA4 in addition to RA1 in 2004, when the cumulative loss of roost trees in RA1 over 3 consecutive years reached 18%. Most bats exclusively roosted in RA4 in 2007, when the cumulative loss of roost trees over 6 consecutive years had reached 46% in RA1. Annual survival for roost trees, from when we first discovered bats using them, was generally lower in RA1 than in RA4. Our results suggest that the movement of bats to the new roosting area corresponded with high losses of roost trees in RA1. This provides additional evidence that to maintain high densities of suitable roost trees for bats in northern temperature forests over several decades, management plans need to recruit live and dead trees in multiple age classes and stages of decay that will be suitable for the formation of new cavities. © 2019 The Wildlife Society.     

Brazilian sleeping beauty: First evidence of hibernation by a bat in South America

Torpor and hibernation are energy-saving heterothermic strategies widespread among mammals, and bats have reports of hibernation in North America, Europe, Asia, Oceania, and Africa. However, to date, there was no evidence of hibernation by bats in South America. We tagged four individuals of Eptesicus brasiliensis and one remained torpid for 13 consecutive days in a tree cavity roost in southern Brazil, showing evidence that bats can hibernate in South America.     

A novel method for targeting survey effort to identify new bat roosts using habitat suitability modelling

In the UK, four out of 18 bat species are listed on the EU Habitats Directive, including the lesser horseshoe bat (Rhinolophus hipposideros), and their population status is closely monitored by visiting known roosts. R. hipposideros predominantly form maternity roosts in buildings, but roosts are impermanent features in the landscape and their distribution changes as bats form new roosts and abandon others. Locating new roosts requires intensive surveys which are challenging and inefficient. In this study, we provide a novel model-based strategy to identify potential R. hipposideros maternity roost sites that can be used to monitor bat populations. First, we model potential maternity roost habitat using record centre data on roost locations across Wales, Great Britain. We then constrain the area identified from modelling using record centre data on locations of bats in areas with no known roosts. We used two variable selection methods and three pseudo-absence data sets (random background points, random points in buildings and target group selection of mammal records) to produce six habitat suitability models. The three pseudo-absence data sets produced different habitat suitability maps, demonstrating the influence of pseudo-absence selection on species distribution models. The six models were combined using weighted mean average to produce an ensemble model that performed better than individual models and that indicated high levels of congruence in areas predicted to have high habitat suitability for maternity roosts. Our model revealed an extensive area (6523 km²; 31% of the area of Wales) containing 18,051 buildings in suitable habitat. Using record centre data on bat activity outside commuting range from known roosts reduced the potential survey area to 133 km² (0.6% of the area of Wales) and 207 buildings. Our modelling outputs can be used to direct volunteers and bat surveyors in more targeted and efficient searches.     

Hibernation by tree-roosting bats

In summer, long-eared bats (Nyctophilus spp.) roost under bark and in tree cavities, where they appear to benefit from diurnal heating of roosts. In contrast, hibernation is thought to require a cool stable temperature, suggesting they should prefer thermally insulated tree cavities during winter. To test this prediction, we quantified the winter thermoregulatory physiology and ecology of hibernating tree-roosting bats, Nyctophilus geoffroyi and N. gouldi in the field. Surprisingly, bats in winter continued to roost under exfoliating bark (65%) on the northern, sunny side of trees and in shallow tree cavities (35%). Despite passive re-warming of torpid bats by 10-20 degrees C per day, torpor bouts lasted up to 15 days, although shorter bouts were also common. Arousals occurred more frequently and subsequent activity lasted longer on warmer nights, suggesting occasional winter foraging. We show that, because periodic arousals coincide with maximum roost temperatures, when costs of rewarming and normothermic thermoregulation are minimal, exposure to a daily temperature cycle could largely reduce energy expenditure during hibernation. Our study provides further evidence that models of torpor patterns and energy expenditure from hibernators in cold temperate climates are not directly applicable in milder climates, where prolonged torpor can be interspersed with more frequent arousals and occasional foraging.     

Roosting behavior and group stability of the big fruit-eating bat Artibeus lituratus (Chiroptera: Phyllostomidae)

The roosting behavior of the big fruit-eating bat, Artibeus lituratus (Phyllostomidae, Stenodermatinae) in an Andean region of Venezuela is described. Sixty-four video recordings made at three separate foliage roosts during 1 year showed that group size varied between two and 14 individuals. One male was regularly observed roosting with more females than others, and this male was associated with the highest quality roost, defined here as the highest, most structurally stable, and least disturbed. Males invariably occupied exclusive roosts during the study, whereas females frequently moved among adjacent roosts. The high roost fidelity of males appears to be related to the defense of the highest quality roosts. Because females were observed roosting with each of the three males present at the study site, female groups were considered unstable. We suggest that shuttling movements of individuals may reflect a commonly observed adaptation of foliage roosting bats related to the avoidance of predators and/or parasites. The type of male–female association observed in A. lituratus is consistent with a resource defense polygyny hypothesis.