Insectivory in captive Livingstone's and Rodrigues fruit bats Pteropus livingstonii and P. rodricensis (Chiroptera: Pteropodidae): a behavioural adaptation for obtaining protein

Captive Pteropus livingstonii and P. rodricensis were observed actively to catch and eat insects. Both species adopt the same technique to trap flying insects, using their wings to envelop them. The spontaneity of the response indicates that it is not an artefact of captivity. The behaviour was observed in captive-bred P. rodricensis which had no prior access to insects, suggesting that it is also innate. A higher rate of successful captures was recorded for an older, wild caught P. livingstonii female, indicating that the effectiveness of the technique used may improve with age. Significant differences in the amount of time spent catching insects by males and females were evident in this species. This study supports suggestions that insects are eaten by some megachiopterans in order to supplement their protein intake.     

Collection, seminal characteristics and chilled storage of spermatozoa from three species of free-range flying fox (Pteropus spp.)

This study reports observations on the collection and characteristics of semen from free-range populations of flying fox in Brisbane, Australia. Semen was successfully recovered by electroejaculation from 107 of 115 wild flying foxes (Pteropus alecto, Pteropus poliocephalus and Pteropus scapulatus). A proportion of ejaculates collected from all three species contained seminal vesicle secretions, the incidence of which appeared related to breeding season. Ejaculate volume was small (5--160 microL), requiring a specialised collection vessel and immediate extension to avoid desiccation. Sperm morphological abnormalities and characteristics are described for the first time. In two species (P. scapulatus and P. alecto), sperm quality varied with breeding season. Dilution in Tris-citrate-fructose buffer and subsequent incubation (37 degrees C) of Pteropus semen for 2-3h appeared to have a negative impact on sperm motility and the percentage of sperm with intact plasma membranes and acrosomes and represents a concern for the potential development and use of assisted breeding technology in these species. Preliminary attempts to develop a short-term chilled preservation protocol for flying fox semen revealed that sperm viability (percentage motility and percentage live sperm with intact acrosomes) was significantly reduced after 102 h chilled storage at 5 degrees C; nevertheless, approximately 40% of the spermatozoa were still motile and contained intact acrosomes. Glycerol was neither protective nor detrimental to sperm survival during chilled storage. Microbial flora of the prepuce, urethra and semen of all species were isolated and their antibiotic susceptibility tested. Tetracycline, penicillin, ciprofloxacin, and ceftazidime were the most effective antibiotics in preventing growth of all identified bacteria; however, their effects on sperm survival were not investigated.     

Abundance, biology, and human exploitation of bats in the Palau Islands

Three species of bats, Pteropus pilosus, P. muriannus pelewensis , and Embullonura semicaudata palauensis , were surveyed by observations, systematic counts, and interviews with local residents in the Palau Islands of Micronesia during a one-month period in 1991. Pteropus pilosus is presumed extinct, with no observations made this century. Pteropus m. pelewensis occurred throughout the archipelago. It is less colonial than other subspecies of P. muriannus , with most of the population appearing to roost solitarily or in small groups of a few bats. Colonies were uncommon and typically ranged in size from about 30–200 animals. Evidence of two larger roosting concentrations, one of which may have held as many as 3,000 bats, was also found. These fruit bats forage and roost in a variety of forested habitats and have diverse diets known to contain the fruits of 54 species, the flowers of 28 species, and the leaves of one species. At dusk, most bats began foraging within several kilometres of their day roosts, but some flew at least 12 km to reach feeding sites. The largest numbers of P. m. pelewensis roosted on rugged and fairly inaccessible limestone islands near the centre of the archipelago, near where most people also lived. We were unable to estimate population size or determine population trend in the island chain. The bats are heavily harvested for commercial export, with more than 180,000 animals shipped to the Mariana Islands from 1975–1994. Smaller numbers are also killed as a local food source. Emballonura s. palauensis , the only insectivorous bat in the islands, was seen at 65% of the evening count locations and was also widespread in the archipelago. Recommendations for managing bats in Palau include a prohibition on the export of fruit bats and the establishment of several protected areas.     

Taxonomy,distribution, and natural history of flying foxes (Chiroptera,Pteropodidae) in the Mortlock Islands and Chuuk State,Caroline Islands

The taxonomy, biology, and population status of flying foxes (Pteropus spp.) remain little investigated in the Caroline Islands, Micronesia, where multiple endemic taxa occur. Our study evaluated the taxonomic relationships between the flying foxes of the Mortlock Islands (a subgroup of the Carolines) and two closely related taxa from elsewhere in the region, and involved the first ever field study of the Mortlock population. Through a review of historical literature, the name Pteropus pelagicus Kittlitz, 1836 is resurrected to replace the prevailing but younger name Pteropus phaeocephalus Thomas, 1882 for the flying fox of the Mortlocks. On the basis of cranial and external morphological comparisons, Pteropus pelagicus is united taxonomically with Pteropus insularis “Hombron and Jacquinot, 1842” (with authority herein emended to Jacquinot and Pucheran 1853), and the two formerly monotypic species are now treated as subspecies — Pteropus pelagicus pelagicus in the Mortlocks, and Pteropus phaeocephalus insularis on the islands of Chuuk Lagoon and Namonuito Atoll. The closest relative of Pteropus pelagicus is Pteropus tokudae Tate, 1934, of Guam, which is best regarded as a distinct species. Pteropus pelagicus pelagicus is the only known resident bat in the Mortlock Islands, a chain of more than 100 atoll islands with a total land area of <12 km². Based on field observations in 2004, we estimated a population size of 925–1,200 bats, most of which occurred on Satawan and Lukunor Atolls, the two largest and southernmost atolls in the chain. Bats were absent on Nama Island and possibly extirpated from Losap Atoll in the northern Mortlocks. Resident Mortlockese indicated bats were more common in the past, but that the population generally has remained stable in recent years. Most Pteropus phaeocephalus pelagicus roosted alone or in groups of 5–10 bats; a roost of 27 was the largest noted. Diet is comprised of at least eight plant species, with breadfruit (Artocarpus spp.) being a preferred food. Records of females with young (April, July) and pregnant females (July) suggest an extended breeding season. Pteropus pelagicus pelagicus appears most threatened by the prospect of sea level rise associated with global climate change, which has the potential to submerge or reduce the size of atolls in the Mortlocks. Occasional severe typhoons probably temporarily reduce populations on heavily damaged atolls, but hunting and ongoing habitat loss are not current problems for the subspecies.     

Interspecific and intraspecific variation in proximate, mineral, and fatty acid composition of milk in Old World fruit bats (Chiroptera: Pteropodidae)

We examine the effect of body mass on milk composition among Old World fruit bats, including Pteropus pumilus (0.175 kg), Pteropus rodricensus (0.265 kg), Pteropus hypomelanus (0.571 kg), and Pteropus vampyrus (1.133 kg). We describe intra- and interspecific differences in the proximate composition of milk among these four species and the minerals and fatty acids in the milk of the latter two species. There were no differences between species in the concentrations of dry matter, fat, or lactose in milk. However, there were significant, although small, differences in the protein content of milk among species, with protein being significantly greater in P. rodricensus than in P. pumilus and P. hypomelanus and protein being significantly less in P. hypomelanus than in P. rodricensus and P. vampyrus. There were no differences in mineral content between P. hypomelanus and P. vampyrus in milk minerals, but minor differences were evident in fatty acids 12:0, 14:0, 18:0, 18:1n11, and 18:2n6. Our findings suggest that milk composition is relatively constant across lactation for most proximate, mineral, and fatty acid components. We found a significant increase in dry matter and energy across lactation in the concentration of dry matter and energy in P. pumilus and fat in P. hypomelanus. In P. hypomelanus, we found a significant increase in the concentration of fatty acids 10:0 and 20:1n9 and a significant decrease in Iso15 and 20:1n7. No other differences associated with day of lactation were found. These findings suggest that milk composition is generally similar within the genus Pteropus, despite a 6.5-fold difference in body mass between species that we evaluated.     

Climate change and the effects of temperature extremes on Australian flying-foxes

Little is known about the effects of temperature extremes on natural systems. This is of increasing concern now that climate models predict dramatic increases in the intensity, duration and frequency of such extremes. Here we examine the effects of temperature extremes on behaviour and demography of vulnerable wild flying-foxes (Pteropus spp.). On 12 January 2002 in New South Wales, Australia, temperatures exceeding 42 degrees C killed over 3500 individuals in nine mixed-species colonies. In one colony, we recorded a predictable sequence of thermoregulatory behaviours (wing-fanning, shade-seeking, panting and saliva-spreading, respectively) and witnessed how 5-6% of bats died from hyperthermia. Mortality was greater among the tropical black flying-fox, Pteropus alecto (10-13%) than the temperate grey-headed flying-fox, Pteropus poliocephalus (less than 1%), and young and adult females were more affected than adult males (young, 23-49%; females, 10-15%; males, less than 3%). Since 1994, over 30000 flying-foxes (including at least 24500 P. poliocephalus) were killed during 19 similar events. Although P. alecto was relatively less affected, it is currently expanding its range into the more variable temperature envelope of P. poliocephalus, which increases the likelihood of die-offs occurring in this species. Temperature extremes are important additional threats to Australian flying-foxes and the ecosystem services they provide, and we recommend close monitoring of colonies where temperatures exceeding 42.0 degrees C are predicted. The effects of temperature extremes on flying-foxes highlight the complex implications of climate change for behaviour, demography and species survival.     

The use of radio-tracking in studying the foraging behaviour of the Indian Flying fox (Pteropus giganteus)

Although much is known about the behaviour of the Indian Flying fox (Pteropus giganteus Brunnich) whilst at the roost (e.g. Neuweiler, 1969), very little is known of its foraging behaviour. The purpose of this paper is to describe the techniques and results of a preliminary radio-tracking study of P. giganteus. In view of the fact that some species of Pteropus are commercial pests (Ratcliffe, 1932) and others are declining rapidly in numbers so as to cause alarm to conservation bodies (Nicoll & Racey, 1981), it is likely that this genus will receive considerable attention in the future. P. giganteus is the largest species of frugivorous bat in Ceylon and is most widely distributed throughout the wet zone (Eisenberg & McKay, 1970).     

Fit females and fat polygynous males: seasonal body mass changes in the grey-headed flying fox

When females and males differ in their timing of maximum reproductive effort, this can result in sex-specific seasonal cycles in body mass. Such cycles are undoubtedly under strong selection, particularly in bats, where they affect flying ability. Flying foxes (Old World fruit bats, Pteropus spp.) are the largest mammals that can sustain powered flight and therefore face critical trade-offs in managing body reserves for reproduction, yet little is known about body mass dynamics in this group. I investigated body mass changes in relation to reproductive behaviour in a large colony of grey-headed flying foxes (Pteropus poliocephalus). In this polygynous mammal, females were predicted to maximise reproductive effort during lactation and males during the breeding season. As predicted, female body condition declined during the nursing period, but did not vary in relation to sexual activity. By contrast, males accumulated body reserves prior to the breeding season, but subsequently lost over 20% of their body mass on territory defence and courtship, and lost foraging opportunities as they also defended their day roost territories at night. Males in better condition had larger testes, particularly during territory establishment, prior to maximum sexual activity. Thus, the seasonality of female mass reflected the high metabolic load that lactation imposes on mothers. However, male mass followed a pattern akin to the "fatted male phenomenon", which is commonly observed in large polygynous mammals with seasonal reproduction, but not in bats. This shows the importance of body reserves for reproduction in flying foxes, despite their severe constraints on body mass.     

Isolation and characterisation of polymorphic microsatellite loci in the vulnerable spectacled flying fox, Pteropus conspicillatus

The spectacled flying fox, Pteropus conspicillatus, is listed as vulnerable in Australia and is under threat from numerous impacts. Primers to amplify eight co-dominant microsatellite loci were designed for Pteropus conspicillatus, based on an enriched genomic library. Four loci were monomorphic in this species while the remaining four loci were highly polymorphic with 16–23 alleles. Two of the four monomorphic loci were found to be polymorphic in Pteropus alecto, a closely related congener. All but one of the six polymorphic loci were in Hardy Weinberg equilibrium. Additionally, six microsatellite loci isolated for Pteropus rodricensis were tested against individuals of P. conspicillatus with all loci amplifying reliably. These loci will be used to investigate population genetic structure in the vulnerable spectacled flying fox.     

Measuring Physiological Stress in Australian Flying-Fox Populations

Flying-foxes (pteropid bats) are the natural host of Hendra virus, a recently emerged zoonotic virus responsible for mortality or morbidity in horses and humans in Australia since 1994. Previous studies have suggested physiological and ecological risk factors for infection in flying-foxes, including physiological stress. However, little work has been done measuring and interpreting stress hormones in flying-foxes. Over a 12-month period, we collected pooled urine samples from underneath roosting flying-foxes, and urine and blood samples from captured individuals. Urine and plasma samples were assayed for cortisol using a commercially available enzyme immunoassay. We demonstrated a typical post-capture stress response in flying-foxes, established urine specific gravity as an attractive alternative to creatinine to correct urine concentration, and established population-level urinary cortisol ranges (and geometric means) for the four Australian species: Pteropus alecto 0.5–305.1 ng/mL (20.1 ng/mL); Pteropus conspicillatus 0.3–370.9 ng/mL (18.9 ng/mL); Pteropus poliocephalus 0.3–311.3 ng/mL (10.1 ng/mL); Pteropus scapulatus 5.2–205.4 ng/mL (40.7 ng/mL). Geometric means differed significantly except for P. alecto and P. conspicillatus. Our approach is methodologically robust, and has application both as a research or clinical tool for flying-foxes, and for other free-living colonial wildlife species.     

Recent loss of vitamin C biosynthesis ability in bats

The traditional assumption that bats cannot synthesize vitamin C (Vc) has been challenged recently. We have previously shown that two Old World bat species (Rousettus leschenaultii and Hipposideros armiger) have functional L-gulonolactone oxidase (GULO), an enzyme that catalyzes the last step of Vc biosynthesis de novo. Given the uncertainties surrounding when and how bats lost GULO function, exploration of gene evolutionary patterns is needed. We therefore sequenced GULO genes from 16 bat species in 5 families, aiming to establish their evolutionary histories. In five cases we identified pseudogenes for the first time, including two cases in the genus Pteropus (P. pumilus and P. conspicillatus) and three in family Hipposideridae (Coelops frithi, Hipposideros speoris, and H. bicolor). Evolutionary analysis shows that the Pteropus clade has the highest ω ratio and has been subjected to relaxed selection for less than 3 million years. Purifying selection acting on the pseudogenized GULO genes of roundleaf bats (family Hipposideridae) suggests they have lost the ability to synthesize Vc recently. Limited mutations in the reconstructed GULO sequence of the ancestor of all bats contrasts with the many mutations in the ancestral sequence of recently emerged Pteropus bats. We identified at least five mutational steps that were then related to clade origination times. Together, our results suggest that bats lost the ability to biosynthesize vitamin C recently by exhibiting stepwise mutation patterns during GULO evolution that can ultimately lead to pseudogenization.     

Flying-fox (Pteropus spp.) sperm membrane fatty acid composition,its relationship to cold shock injury and implications for cryopreservation success

The very large acrosome of Pteropus species spermatozoa is prone to damage during cooling procedures. Cryogenic succuss has been linked to membrane composition, therefore the lipid composition of five Pteropus species sperm acrosomal and plasma membranes were investigated to provide insight into reasons for cold shock susceptibility. Rapid chilling and re-warming of spermatozoa from three Pteropus species resulted in a decrease (P < 0.05) in acrosomal integrity. Biochemical analysis of lipids revealed that stearic acid (18:0) was the predominant saturated fatty acid and oleic acid (18:1, n-9) the predominant unsaturated fatty acid in both acrosomal and plasma membranes. Linolenic acid (18:3, n-3) was only detected in plasma membranes of Pteropus hypomelanus and was detected in acrosomal membranes of all Pteropus spp. studied (except Pteropus giganteus). Although detected in both plasma and acrosomal membranes of Pteropus vampyrus, docosahexaenoic acid (22:6) was not detected at all in Pteropus poliocephalus, only in trace levels in the acrosomal and plasma membranes of P. giganteus and P. hypomelanus and not in acrosomal membranes of Pteropus rodricensis. No difference was seen in the levels of polyunsaturated fatty acids (PUFAs) within plasma membranes, however PUFAs were lower (P < 0.05) in acrosomal membranes of P. giganteus compared with P. vampyrus. Pteropus spp. spermatozoa have a very low ratio of unsaturated/saturated membrane fatty acids (<0.5). Membranes containing more PUFAs are more fluid, so the use of cryogenic media which improves membrane fluidity should improve Pteropus spp. spermatozoal viability post-thaw.     

Depth equilibration by two predatory cichlid fish from Lake Malawi

The cichlid fish, Haplochromis polystigma and Haplochromis livingstonii are piscivorous ambush predators endemic to Lake Malawi, Africa. Experimentally it was found that large adult H. livingstonii could equilibrate to 38 m, but that juveniles were restricted to shallower depths. Mean depth attained by 8 H. polystigma was 45 m but two individuals exceeded the maximum simulated depth (59 m). H. livingstonii had a mean equilibration rate of 4–7 m day⁻¹ for the first 10 m declining to 2·4 m day⁻¹ thereafter. The mean rate of descent for H. polystigma was 3·8 m day⁻¹ for the first 10 m and 2·8 m day⁻¹ thereafter. The fish were decompressed at a rate of a 20% decrease every 12 h. Although cichlid fishes occur down to 200 m depth in Lake Malawi, experimental and field data indicate that the depth distribution of individual species is restricted and that the restriction is largely related to swimbladder physiology. All cichlid species so far tested equilibrate slowly to depth (2 to 5 m a day) and are not capable of rapid decompression.     

Flying-fox (Pteropus spp.) sperm membrane fatty acid composition,its relationship to cold shock injury and implications for cryopreservation success

The very large acrosome of Pteropus species spermatozoa is prone to damage during cooling procedures. Cryogenic succuss has been linked to membrane composition, therefore the lipid composition of five Pteropus species sperm acrosomal and plasma membranes were investigated to provide insight into reasons for cold shock susceptibility. Rapid chilling and re-warming of spermatozoa from three Pteropus species resulted in a decrease (P < 0.05) in acrosomal integrity. Biochemical analysis of lipids revealed that stearic acid (18:0) was the predominant saturated fatty acid and oleic acid (18:1, n-9) the predominant unsaturated fatty acid in both acrosomal and plasma membranes. Linolenic acid (18:3, n-3) was only detected in plasma membranes of Pteropus hypomelanus and was detected in acrosomal membranes of all Pteropus spp. studied (except Pteropus giganteus). Although detected in both plasma and acrosomal membranes of Pteropus vampyrus, docosahexaenoic acid (22:6) was not detected at all in Pteropus poliocephalus, only in trace levels in the acrosomal and plasma membranes of P. giganteus and P. hypomelanus and not in acrosomal membranes of Pteropus rodricensis. No difference was seen in the levels of polyunsaturated fatty acids (PUFAs) within plasma membranes, however PUFAs were lower (P < 0.05) in acrosomal membranes of P. giganteus compared with P. vampyrus. Pteropus spp. spermatozoa have a very low ratio of unsaturated/saturated membrane fatty acids (<0.5). Membranes containing more PUFAs are more fluid, so the use of cryogenic media which improves membrane fluidity should improve Pteropus spp. spermatozoal viability post-thaw.     

Bats Without Borders: Long-Distance Movements and Implications for Disease Risk Management

Fruit bats of the genus Pteropus (commonly known as flying-foxes) are the natural hosts of several recently emerged zoonotic viruses of animal and human health significance in Australia and Asia, including Hendra and Nipah viruses. Satellite telemetry was used on nine flying-foxes of three species (Pteropus alecto n = 5, P. vampyrus n = 2, and P. neohibernicus n = 2) to determine the scale and pattern of their long-distance movements and their potential to transfer these viruses between countries in the region. The animals were captured and released from six different locations in Australia, Papua New Guinea, Indonesia, and Timor-Leste. Their movements were recorded for a median of 120 (range, 47–342) days with a median total distance travelled of 393 (range, 76–3011) km per individual. Pteropus alecto individuals were observed to move between Australia and Papua New Guinea (Western Province) on four occasions, between Papua New Guinea (Western Province) and Indonesia (Papua) on ten occasions, and to traverse Torres Strait on two occasions. Pteropus vampyrus was observed to move between Timor-Leste and Indonesia (West Timor) on one occasion. These findings expand upon the current literature on the potential for transfer of zoonotic viruses by flying-foxes between countries and have implications for disease risk management and for the conservation management of flying-fox populations in Australia, New Guinea, and the Lesser Sunda Islands.     

Flying foxes as carriers of pathogenic Leptospira species

Recent serologic studies have identified flying foxes (Pteropus spp.) as carriers of leptospirosis; however, little is known about the role of flying foxes as carriers of pathogenic Leptospira spp. To determine if Australian Pteropus spp. are carriers of pathogenic Leptospira spp., TaqMan real-time polymerase chain reaction (PCR) was used to detect leptospiral DNA in kidney and urine specimens from four species of flying fox, including the spectacled flying fox (Pteropus conspicillatus), black flying fox (Pteropus alecto), grey-headed flying fox (Pteropus poliocephalus), and little red flying fox (Pteropus scapulatus). Of the 173 kidney samples tested, 19 (11%) were positive for leptospiral DNA. Positive individuals were detected in all four species; significant differences in prevalence were not detected between species, between species within the same geographic area, or between geographically separated samples from the same species. Of the 46 urine samples tested, 18 (39%) tested positive by PCR, confirming that flying foxes shed leptospires into the environment. The detection of leptospiral DNA in the kidneys and urine of flying foxes suggests that flying foxes are carriers of pathogenic Leptospira spp. No evidence collected in the present study, however, suggests that flying foxes pose a significant risk of leptospirosis to the wider community or that humans who are in regular, close contact with flying foxes are at risk for leptospirosis.     

Taxonomic Studies of Phallodrilinae (Oligochaeta, Tubificidae) from the Great Barrier Reef and the Comoro Islands with Descriptions of Ten New Species and One New Genus

A taxonomic and morphological account of a collection of Phallodrilinae from coralline sands at Heron Island in Australia's Great Barrier Reef is given. Ten new forms are described: Jamiesoniella athecata gen. et sp.n. Phallodrilus rectisetosus heronensis subsp.n. P. geniculatus sp.n. P. filithecatus sp.n. P. clavatus sp.n. (latter three all devoid of an alimentary canal), Bathydrilus superiovasatus sp.n. Coralliodrilus atriobifidus sp.n. C. oviatriatus sp.n. C. parvigenitalis sp.n. and the gutless C. avisceralis sp.n. Morphological notes are given for Phallodrilus albidus Jamieson, 1977 and Bathydrilus rohdei (Jamieson, 1977). The new meiobenthic genus Jamiesoniella is characterized by having simple atria with only one pair of prostate glands, which are attached to ectal parts of atria, and by lacking spermathecae and penial setae, The genus is probably related to Aktedrilus Knöllner and Bacescuella Hrabě. A gutless species, Phallodrilus comorensis sp.n. is described from the Comoro Islands in the Mozambique Channel (W Indian Ocean).     

Allomaternal care: helper-assisted birth in the Rodrigues fruit bat, Pteropus rodricensis (Chiroptera: Pteropodidae)

We report observations of allomaternal care, or epimeletic (care-giving) behaviour, exhibited by one female toward another, before, during, and after giving birth, in the Rodrigues fruit bat, Pteropus rodricensis (Chiroptera: Pteropodidae). Interactions between the expectant mother and other roost mates, from the onset to completion of birth, were observed continuously over a period of approximately three hours. The attending female assisted the mother when she: (I) intermittently groomed her anovaginal region; (2) grasped her with partially outstretched wings; (3) 'tutored' her in a feet-down birthing posture; (4) groomed the emerging pup; and (5) physically assisted the mother by manoeuvring the pup into a suckling position. This is the first report of extensive assistance given by a female bat to another during parturition. We suggest that epimeletic behaviour may be common in the highly social Chiroptera, especially among primiparous or inexperienced females and/or when an expectant mother experiences difficulty during labour and parturition. Our observations support the hypothesis that roosting associations in bats may be as important as relatedness in explaining the occurrence of apparently altruistic acts.     

The hormonal and behavioral response to group formation, seasonal changes, and restraint stress in the highly social Malayan Flying Fox (Pteropus vampyrus) and the less social Little Golden-mantled Flying Fox (Pteropus pumilus) (Chiroptera: Pteropodidae)

This study examined behavioral and physiological responses (changes in inter-animal spacing, total glucocorticoids, testosterone, and body mass) to the formation of breeding and same-sex groups in two bat species, the socially gregarious Malayan Flying Fox (Pteropus vampyrus) and the less social Little Golden-mantled Flying Fox (Pteropus pumilus). We hypothesized that social instability, especially in the breeding groups and especially in P. vampyrus, would result in elevated glucocorticoids and that social facilitation of breeding and/or male-male competition would result in persistently higher levels of testosterone in breeding males. Seasonal rhythms in all measures were also predicted, and the glucocorticoid stress response was expected to vary by sex, season, and group type. Nearly all animals responded to group formation with elevated glucocorticoids, but, for breeding animals (especially aggressive male P. vampyrus), these responses persisted over time. In both species, breeding group formation resulted in elevated testosterone in males. Glucocorticoids, testosterone, testes volume, and body mass generally peaked in the breeding season in males (late summer and early autumn), but the seasonal glucocorticoid peak in females occurred in late winter and early spring. All animals responded to restraint stress with elevations in glucocorticoids that largely did not differ by sex, time of year, reproductive condition, group type, or, in lactating females, the presence of her pup. Changes in both behavior and physiology were more evident in P. vampyrus than in P. pumilus, and we believe that their underlying social differences influenced their responses to group formation and to the changing seasonal environment.     

Further examination of the basicranial anatomy of the megachiroptera: a reply to A.J. King.

Four basicranial characters supporting a single origin of megachiropterans and microchiropterans have been identified by Wible and Novacek. King was not able to find two of these characters in specimens of the megachiropteran Pteropus and concluded that these characters be rejected in analyses of chiropteran phylogeny. To address King's claims, the two characters in question (regarding the tegmen tympani and ramus inferior of the stapedial artery) are documented here for Pteropus. The discrepancies between this report and King's appear to result from different criteria for identifying the tegmen tympani. Employed here are the criteria generally accepted by students of the mammalian chondrocranium.