Avian prey of the Australian Ghost gigas (Microchiroptera: ): prey characteristics and damage from predation

Walter E. Boles1.' 'Division of Vertebrate Zoology, Australian Museum, 6 College Street, Sydney, New South Wales 2000 2School of Biological Science, Univeriity of New South Wales, New South Wales 2052

ABSTRACT The Macroderma gigas is a large (mean mass 150 g) predatory bat of sub- tropical and tropical . It carries its vertebrate prey to roost caves to be eaten and where remains are dropped and accumulate. Whereas the attack and feeding methods of M. gigas on has been well documented, there is little comparable information relating to avian prey. Using published lists of avian prey and skeletal material collected from a modern Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021 Macroderma roost, this study examined the range of bird species eaten by this bat. From the characteristics of these birds, biases towards particular behaviour patterns were identified. Prey masses were used to determine a preferred size range for avian prey. From this information, and assessments of the damage to the bones, inferences were made regarding the capture and processing methods employed by M. gigas for birds. More than 50 species, from a broad taxonomic range, have been recorded as avian prey of the Ghost Bat. These are all essentially diurnal with the obvious exception of the Australian Owlet-nightjar Aegotheles crisfatus. Species that aggregate when roosting are over-represented in the diet of the Ghost Bat. Ground-frequenting species comprise about a quarter of the prey records. Birds may be captured at most levels of the strata, from the ground to the canopy, and in flight. Ghost feed on a wide size range of avian prey, and although they may take up to about two-thirds their own mass, they have a preference for smaller birds, with almost 70% of the species having a mass less than 35 g. The most severe damage to a bird's post-cranial skeleton is to the ventral side of the sternum, whereas distal appendicular elements exhibit no damage from Ghost Bat predation. The type of damage to the bones and the size of the prey is consistent with the assumption that megadermatid bats were accumulators of many avian remains now represented in and fossil deposits at Riversleigh, Queensland. Key words: Ghost Bat, Bat edict, Avian prey, Fossil bat prey, Macroderma gigas.

INTRODUCTION (Guppy and Coles 1983; Kulzer et al. 1984; Tidemann et al. 1985). Ghost Bats emerge The Ghost Bat Murodema gigas is a large from their roosts and commence hunting (up to 165 g; mean 150 g) predatory bat of about 1 to 1.5 hours after sunset for a period subtropical and tropical Australia (Richards of two hours. This is followed by periods of and Hand 1995) (Fig. 1). It feeds on both inactivity, punctuated by short bursts of vertebrate and invertebrate prey, which are foraging. Resumption of feeding activity takes carried back to roost caves to be eaten, and place in the early morning, up to just before where their remains are dropped to the sunrise (summarized from Tidemann et al. cave floor. Eventually discarded portions 1985). The initial foraging behaviour of accumulate, providing an indication of the M. gigas has been characterized as "sit and prey species of the Ghost Bat. Although inspect", in which bats quietly visually inspect M. gigas is the only species of megadermatid their surroundings for movement from a perch now occurring in Australia, eight Australian (Tidemann et al. 1985). Apparently they are fossil species have been recorded (Hand 1985, unable to detect motionless animals. Once 1995, 1996; Hand et al. 1988). These have prey is located, it may be captured in the been implicated as the source of the accumu- air, gleaned (taken from the surface of the lations of some of the deposits of small substrate by a flying bat) from the ground or vertebrates discovered in the Tertiary fossil foliage, or dropped on from a perch. The bats deposits of Riversleigh, northwestern Queens- are dependent on stealth and darkness to land (Boles 1995, 1997, 1998; Hand 1990, capture prey. Toop (1985) noted that a bird 1995, 1996). in an illuminated room was able to elude capture. Published work on M. gigas has included lists of prey species identified from active Using mice as prey under captive conditions, -~~~~roosts (Douelas 1967: Vestiens and Hall 1977; Kulzer et al. (1984) reported that an attack \ " . . Pettigrew et al. 1985; TOO< 1985; Schulz 1986) flight by M. gigas culminated when the bat and studies of foraging methods, including dropped on its prey, enclosing it with the attack methods on mammals, by this species wings and immediately biting it on the

82 Australian Zoologisf 31(1) June 1999 Pine Creek

FRzroy Caves, Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021

Figure 1. Distribution of Macroderma gigas, showing localities discussed in this paper from which avian prey remains have been identified: (1) several (1 sites in the Pilbara district of Western Australia (Douglas 1967); (2) Pine Creek area, Northern Territory (Pettigrew et al. 1985; Schulz 1986, this study); (3) Fitzroy Caves National Park and Mt Etna in Queensland (Toop 1985). Dark grey is current distribution; light grey indicates area from which species is now extirpated (modified from Richards and Hand 1995). d head, neck or throat. If this did not result in prey of M. gigas have been made almost immediate death, the prey was positioned for exclusively on discarded heads and feathers; additional bites to the head. Prey up to 60 g as yet no author has indicated that identi- was carried to the roost with the head in the fications were based on bones. To date, no bat's jaws. Consumption commenced with the attention has been focused on the manner skull. Smaller mice (c. 10 g) were completely of feeding on and resultant damage by this eaten, whereas parts of adults (remnants of predatory bat species to avian prey. skull, legs, digestive tract and tail) were Thus, using avian skeletal material collected sometimes dropped to the floor. Guppy and Coles (1983) gave similar descriptions of the from a modern M. gigas roost, the type of damage to the bones is described in this feeding behaviour of captive M. gigas. Douglas (1967) noted that the bats ate the whole study. This information is used for indications carcass of mammalian prey, including the fur, of the attack and processing methods that although portions were often dropped. the bats use on avian prey, particularly in comparison with published information on In contrast to several studies that have been mammalian prey. From published lists of conducted on mammalian prey, there is little prey and the remains included in this study, information on how M. gigas attack or process characteristics of prey species are reviewed birds as prey. Pettigrew et al. (1985) observed for any biases towards particular taxonomic a M. gigas unsuccessfully pursue a flying groups, or foraging or other behavioural quail Coturnix (presumably Brown Quail patterns. Avian predators, such as owls, that C. ypsilophora). Douglas (1967) noted that in serve as accumulators, can often be identified captivity the bats grasped dead birds across to species based on the size range of the the chest and recorded that when they ate prey. Using the lists of prey, the size range of birds, "the feathers were discarded where the avian prey of M. gigas is assessed. These possible and wing and tail quills, and the legs, results are also evaluated for consistency with were dropped to the ground". There do not the idea that Miocene and Pliocene Macroderma appear to be any other descriptions of the were responsible for the accumulations of methods of attacking, killing or transporting some of the avian remains found as fossils at avian prey. Published identifications of avian Riversleigh.

Australian Zoologist 31(1) 83 METHODS An exception is Schulz (1986), who estimated Prey species the number of individuals of each taxon of prey at a Ghost Bat roost site, warning that it The list of prey species was compiled from 'has difficult to assess the abundance of prey published records from across the distribution items" and the figures were only approximate. of M. gigas. Sites from which prey records were It is thus not possible for this study to taken include the Pine Creek area, Northern calculate a "mean" prey mass. Territory (Pettigrew et al. 1985; Schulz 1986, this study), Fitzroy Caves National Park and The avian prey species have been tabulated, Mt Etna in Queensland (Toop 1985), and with their masses, in Appendix 1. A histogram several sites in the Pilbara district of Western of these masses in Figure 2 uses the minimum Australia (Douglas 1967) (Fig. 1). values given for each species. It should be heeded that none of the values is quantified Prey size in terms of number of individuals per prey The mass of each avian prey species has species. been taken primarily from Hall (1974), Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021 Marchant and Higgins (1993), Higgins and Prey damage Davies (1996) and specimen records in the The bones used in this study come from the Ornithology Section of the Australian Museum. same general mine complex near Pine Creek Most of the published avian masses are from as that of Pettigrew et al. (1985) and Schulz southern Australian individuals, which are (1986), although not necessarily from the same often larger than conspecifics in the northern mine. Surface material was collected by S. J. part of the country (Bergmann's Rule; Ford Hand, M. Archer, H. Godthelp, K. Aplin and 1989). While this may cause over-estimation of K. Gollan (UNSW) from a series of adits the prey sizes, it does not unduly distort the (horizontal or nearly horizontal passages) in relative distribution of prey species sizes. To Kohinoor Mine, near Pine Creek, conducted compensate for this to some degree, the lower on 10 May, 1983. Preliminary identifications value of the ranges have been used as the were made by G. F. van Tets (CSIRO), a basis for discussion. few of which have been altered followinna Few published studies report the number of subsequent examination by the author. The each prey species present; most merely indicate prey species and skeletal elements recorded the presence of a species in prey remains. are given in Appendix 2. 14 I

Figure 2. Histogram showing range and frequencies of minimum avian prey masses (g) taken from Appendix 1. The indeterminate crake Porzana has been omitted because of the considerable difference in mass between the two species that might be represented.

84 Australian Zoologist 31(1) June 1999 RESULTS appears to be unrelated to predation by Prey species M. gigas. Douglas (1967) identified some avian prey from heads found in roosting caves, hut Over 50 species of birds have been identified did not indicate what sort of damage these as prey of M. gigas (Appendix 1). Twenty-two had incurred. species were recorded from Mt Etna by Toop (1985), (although he named only three Pectoral girdle of these). This compares with 21 species Several coracoids were recovered intact. On identified from the Pilbara (Douglas 1967) and one badly damaged sternum the posterior 40 from Pine Creek (Pettigrew et al. 1985; ends only of the coracoids were still in contact Schulz 1986, this study). Only one species, the and the shafts showed marked laceration Australian Owlet-nightjar Aegotheles cristatus, (Fig. 3A). In two other instances the sterna was recorded from all three localities. A few were badly chewed but the still attached species were found in both the Northern coracoids were undamaged (Fig. 3D). A scapula Territory and Western Australian roosts. Most had minor damage to its posterior tip, which species were represented at only one site. may have been the result of chewing by Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021 These encompass a broad taxonomic range, M. gigas. but with songbirds (Passeriformes) dominating, comprising about two-thirds of the species. Sternum Most major bird groups are represented with All the four sterna examined had sustained the exceptions of waterbirds (other than severe damage (Fig. 3A-D). The keels were crakes), raptorial birds and those containing heavily lacerated along their ventral margins, only very large-bodied forms. All species were showing extensive evidence of chewing, and in essentially diurnal with the exception of several instances this extended through the Aegotheles crktatus. A substantial number of the entire depth of the keel to the sternal plate. prey species are ground-frequenting (n = 19) The damage also occurred along at least one or aggregate when roosting (n = 15). of lateral sides of the sternum. In some Among the avian remains examined in this instances the sternal plate had been removed study were several that represented young almost to the base of the keel. The posterior birds, as indicated by the pitted surface of portion of the sternum was missing in all the bone and incomplete ossification of the instances. articular facets (for example, cf. Silver-crowned Friarbird Philemon argenticeps; Appendix 2; Fig. 3H). No complete pelvic elements were found. In one specimen, the anterior end of the ilium Prey size appeared to have been chewed to about the Throughout the distribution of M. gigas, acetabulum; the ischium and pubis showed no prey species in the size range of 6-40g such damage (Fig. 3G). The anterior vertebrae predominate, but in the Northern Territory of the synsacrum (thoracic, lumbar) were population this upper boundary extends to not recovered. Two sets of sacral vertebrae 50-60 g. Very few heavier birds (>70 g) have were detached from the pelvic elements and been recorded. These assessments are based appeared to have been chewed on the anterior on the species list without allowance for the end and for part of the lateral edge on one numbers of individuals in each size class. side (Fig. 3E-F). Schulz's (1986) rough estimates indicated that Wing and leg within this broad size range there was a bias towards the smaller end of the scale, with Distal elements of wings (ulna, radius, about a third of the prey items falling in the carpometacarpus, digits) and legs (tibiotarsus, 10-20 g class. Of the prey species in Appendix tarsometatarsus) exhibited no indication of 1, almost 70% have a mass of 35 g or less damage from M. gigas. No humeri were (Fig. 2). recovered except for an instance in which the wing, sternum and coracoids remained joined. Prey damage (Fig. 3) In this case, most of the humerus was missing, with only the extreme proximal and distal Skull ends being present, held together and Only one avian skull was examined. It attached to the remaining bones by dried showed minor damage to the rear, which may tissue (Fig. 3B). The loss of the shaft may have not have resulted from M. gigas predation. resulted from chewing. The only femur There was no injury of the magnitude recovered was a distal-most iragment; the described for the consumption of mice. A slight damage to it did not appear to be due bill of a kingfisher exhibits breakage that to M. gigas.

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Figure 3. Avian bones from Kohinoor Mine, Northern Territory, damaged by Ghost Bats. A. Sternum of Trichoglossus haematodus with remnant of coracoid. B. Sternum, pectoral girdle and forelimb of Geopelia cuneaia, showing proximal fragment of humerus. C. Sternum of Geopelia striata. D. Sternum of %mix castanota with undamaged coracoids. E. Synsacrum of Geopelia striata. F. Synsacrum of Trichoglossus haematodus. G. Pelvic fragment of Geopelia cuneata. H. Tibiotarsus of large honeyeater (cf. Phibmon argenticeps) showing porous surface and incompletely formed articular surfaces, indicative of immaturity. I. Tibiotarsus of Philemon argenticeps exhibiting post-mortem damage to shaft not resulting from Ghost Bat predation. Bar equals 10 mm for all figures except G, for which it equals 5 mm.

DISCUSSION tend to aggregate when roosting. The most obvious of these are the four species of Prey species woodswallows Artamus. Individuals often roost Based on the list of recorded avian prey, in close proximity, a habit that presumably M. gigas appears to select its prey oppor- makes them more easily located. Other tunistically. Among the wide range of social species that roost in close aggregations taxonomic groups, the only ones that are include the lorikeets Trichoglossus and over-represented are those with species that Psitteuteles, Budgerigar Melopsittacus undulatus,

86 Australian Zoologist 31 (1) June 1999 Rainbow Bee-eater Merops ornatus, Varied they were captured on the ground (note Sittella Daphoenositta chrysoptera and finches observation by Pettigrew et al. (1985) of the Taeniopygia, Poephila, Erythura and Emblema. attempted capture of a flying quail). Many of These are indicated in Appendix 1 and Figure the birds that nest and forage on the ground 2. Individuals of the Grey-crowned Babbler roost above it, as do almost all of the other Pomatostomus temforalis roost in close associa- recorded prey species. The range of avian prey tion but within an enclosed shelter. species indicates M. gigas will capture birds at The nocturnal Australian Owlet-nightjar almost any level, from the ground or elevated Aegotheles cristatus is the obvious exception perches, and in flight. among the otherwise primarily diurnal avian It is known that inexperienced young birds prey species. It emerges and becomes active at have a higher mortality from predators than the same time as M. gigas. Because it could be do older individuals (Newton 1993; Sullivan captured not only when perched, but also in and Roper 1996). and it is likely that flight or on the ground when feeding, it is young birds form a significant proportion of not surprising that this species was recorded the prey of M. gigas. They would be more by four studies. Other species, such as quail easily captured and possibly less skilled in Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021 Coturnix, button-quail Turnix and crakes defending themselves, thus reducing potential Porzana, are frequently active at night, and injury to the bat. This could be particularly a number of species vocalize on moonlit important with larger birds with strong bills nights; both these behaviours would increase and claws, such as the young Pied Butcherbird the risk of predation by M. gigas. Most diurnal Cracticw nigrogularis recorded by Schulz (1986). birds will have gone to their roosts for the Old or weak birds might also be expected night before the bats emerge, but may not to form a disproportionate amount of the have completely settled. Any movement would prey; however, these are features that cannot make them potentially detectable to the bats. be assessed from examination of the remains. Many species begin vocalizing and becoming The list of prey species shows differences active before sunrise, corresponding with the between the colonies of M. gigus, as well as early morning foraging bout of the bats, again occurrences common to two or all colonies making them vulnerable to detection and (Appendix 1). In some cases, a widespread attack. bird species that occurs throughout the At a central coastal Queensland colony, Toop distribution of M. gigas bas been recorded as (1985) noted that ground-dwelling prey were prey from only one location (e.g., Merops rarely taken, and contrasted this with the ornatus). This is likely to be an artefact of the practice of M. gigas in desert habitats. The small number of samples. Other prey species three species named of the 22 he recorded live in only part of the predator's range. The were arboreal. The lists of prey species from few records from the Mt Etna colonies drier country colonies of M. gigas of the in Queensland include several rainforest Northern Territory and Western Australia species (e.g., Melzphaga lewinii, Colluricincla confirm that a substantial proportion is ground- megarhyncha), whereas among the avian prey of frequenting. A somewhat arbitrary delimitation the Pilbara colonies are desert species (e.g., of ground-frequenting species comprises those Amytornis striatus, Cinclosoma castaneothorax). that feed and nest on the ground (quail There are insufficient published records from Cotumix, button-quail Turnix, crake Porzana, the coastal eastern colony to make further Spinifex Pigeon Geophaps plumifera, Chestnut- comparisons. breasted Quail-thrush Cinclosoma castanaotho~ax, Rufous Songlark Cincloramphus mathewsi, Prey size Richard's Pipit Anthus nouaeseelandiae) and The records compiled here indicate that those feed extensively on the ground but roost M. gigas will take a range of avian prey sizes and nest in trees or shrubs (doves Geopelia, but generally selects smaller species, with a Red-backed Fairy-wren Malurus melanocephalus, third of those recorded falling below 20 g and Striated Grasswren Amytornis striatus, Grey- more than two-thirds less than 35 g. The crowned Babbler Pomatostomus and finches extent to which this choice of prey species Taeniopygia, Poephila, Erythura and Emblema). is actually based on size, rather than factors These groups are designated in Appendix 1 such as opportunism, prey detectability and and Figure 2. Seven of the species made roosting habits cannot be determined. The up about one-third of the individual avian records of large prey species (>80 g) in the remains from 29 species identified by Schulz wild, although few, are interesting. If the (1986). Likewise nine of the 40 species recorded individual birds captured were towards the by Douglas (1967) fall into these categories. high end of their mass range, it would mean That these species are largely ground- that the bats were taking prey about two-thirds frequenting does not necessarily mean that their own mass. As well as the increased

Australian Zoologist 31(1) 87 difficulty of killing such large prey, the chance of the wing (remiges) and tail (rectrices), of injury is increased and ability to return which Douglas (1967) noted are dropped these to the roost for consumption is com- by feeding birds. These feathers are generally promised. Kulzer et al. (1984) reported that the most easily noticed and collected, and captive bats would kill and transport rats up usually among the easiest to identify. The to 80% of their body mass. This may be an remiges are connected to the bones of the artefact of the sizes of food animals offered hand, and so would remain attached to the and probably should be extrapolated to wild discarded wings. situations with care. The record of a Pied Butcherbird Cractzcus nigrogtalaris, a species The small sample of modem bones collected that as an adult would both be heavy and offer in this study agrees with the pattern of the most risk of injury, refers to a young bird small avian bones recovered at Riversleigh (Schulz 1986). which would not present these (Boles 1995, 1997, 1998). At Riversleigh, there problems to the same degree. Kulzer et al. were ample distal elements (carpometacarpi, (1984) noted that captive M. gigas were able tibiotarsi, tarsometatarsi) in the fossil deposits. These were often broken but the damage to kill 100 g young rats without sustaining any Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021 injury. appears to be post-depositional, Coracoids, although much fewer than distal long bones, exceed proximal long bones. Humeri and Prey damage femora from small-bodied forms are unusual. It is not evident from the remains what Skulls, sterna and pelvic elements are even attack methods are employed by M. gigas on less commonly found, no doubt due to their birds; however, there is no reason to believe comparative fragility compared to the long that a novel technique is used relative to those bones. observed with non-avian prey. The patterns of damage to the post-cranial bones of avian prey Conclusiom in relation to Riversleigh remains are consistent with the general observations by Examination of avian prey remains from Kulzer et al. (1984) that mice are consumed modern Ghost Bat colonies has shown that the along the anteroposterior midline of the body. skeletal elements represented and the manner At the same time, remains of sterna suggest of damage these have sustained from M. gigas that M. gigas feeds on birds in a manner predation are consistent with findings on somewhat different from that used for avian fossils from Riversleigh. This supports mammals. In contrast to observations on mice the assumption that megadermatid bats used in feeding experiments (Douglas 1967; were accumulators of avian remains in these Guppy and Coles 1983; Kulzer et al. 1984), deposits. Owls are well documented as birds are not eaten whole. That the major accumulators of vertebrate remains in caves damage is to the keel and lateral edges of in Australia and elsewhere (Baird 1991). To the sternum, while the coracoids frequently date, however, no evidence of owls has been escape any damage, suggests that the principal recovered from the Riversleigh deposits. portion of a carcass consumed is on the ventral side of the bird, presumably the There is also strong concordance between pectoral muscle. The loss of the posterior end the size of the prey animals from the fossil of the sterna in all four of the remains that sites and modern Ghost Bat caves. This agree- were examined suggests that entry may also ment extends to the few taxa that have be made into the body cavity. thus far been identified from the Riversleigh There was no evidence available to assess megadermatid accumulations. Taxa that have been recognized include a small kingfisher whether the skull of birds sustained the same damage as did those of mice in experiments. (Boles 1997), Budgerigar Me1opsdtm-w undulohLs (Boles 1998), and a swallow and honeyeaters If feeding by the bats had commenced at the skull and continued posteriorly, as described (unpublished data). by Kulzer et al. (1984) for mice, then the coracoids would be expected to sustain more ACKNOWLEDGEMENTS damage than was observed. I thank the collectors of the Pine Creek The absence of damage to distal appendicu- material for permitting me to work on it; lar elements is consistent with observations S. J. Hand for information concerning that these portions are discarded by the bat the collection of these specimens and fossil before feeding. Identifications of avian prey by megadermatids, and for comments on the previous authors have been based largely on manuscript; and the late G. E van Tets for feathers, without indicating which feathers his initial identifications. The manuscript were used. It is probable that most of the benefited considerably from comments by D. latter records were based on the large feathers Lunney, S. Burnett and an anonymous referee.

68 Australian Zoologist 31(1) APPENDIX 1 Recorded avian prey species of the Ghost Bat Momhgigas, giving body masses (see text for sources) and references to localities for species' occurrence as prey. (I) Pilbaa, Western Australia (Douglas 1967); (2) Pine Creek, Northern Territory (Pettigrew ct 01. 1985); (3) Pine Creek, Northern Territory (Schulz 1986); (4) Pine Creek, Northern Territory (this study); (5) Mt Etna, Queensland (Toop 1985). ** - feeds and nests on the ground; - feeds but not nests on the mound'; A - amregates when mostina.

Scientific Name English Name ** +Coturnk ?ypsilophoro Brown? Quail ** Ponnna purilloitabucmis Baillon'slSpotless Cnke ** Tumk mnculorn Red-backed Button-quail ** Tumk velox Little Button-quail ** Tumix castanota Chestnut-backed Button-quail Geopclio rhioto Peaceful Dove * Geopelio cuncnln Diamond Dove Gcophnps plumifera Spinifex Pigeon A Trizhoglorsur hnitMIodur Rainbow Larikeet A Psittcuteles virsicolor Varied Loriket Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021 Pscphotus dirrimilis Hooded Parrot A Mclnprittacw undulotur Budgerigar Aegothelrs crirlntus Australian Owlet-nightjar Apw pacifiur Fork-tailed Swift Alcedo arureur Azure Kingfisher Todirnmphus sp. kingfisher [sanctur/pyrrhopygia] Todirnmphm ?macleayh Forest? Kingfisher Todiramphus pyrrhopygia Red-backed Kingfisher A Merops omatur Rainbow Bee-eater Eurysfomur orientalis Dollarbird Climncterir melonura Black-tailed Treecreeper Pardalotvr shiatur Striated Pardalote Gergyone olivncen White-throated Gerygone Malurur melanorebhalus Red-backd Fairy-wen Striated Grasswren Silver-crowned Friarbird " Little Friarbird Monorim flnviguln Yellow-throated Miner Mcliphagn lminii Lewin's Honeyeater Lichenosfomur virescem Singing Honeyeater Lichenostomw keartlnndi Grey-headed Honeyeater Li~henostamusbmicillatus White-plumed Honeyeater Yellow-tinted Honeyeater Brown Honeyeater Melonodryor cucullato Hooded Robin Microcco fnscinons Jacky Winter A Daphocnosittn ihrysopfera Varied Sittella Pachycephola wjivcntris Rufous Whistler ColluricmCln mprhynchn Little Shrik-thrush Colluricioclo harmonica Grey Shrike-thrush Rhipidrrra filiginosa Grey Fantail Rhipidurn nrjvrnhls Northern Fantail ' A Ibnatostomvs temporalis Grey-crowned Babbler ** #Cincloromo cartnncothorox Chestnut-breasted Quail-thrush Lolage sueurii White-winged Triller A A~fonuslevcorynchur White-breasted Woodwallow A Artamw personotus Masked Woodswallow A Artamvr cimmu Black-faced Woodswallow A Artamus minor Little Woodswallow Croctirw nigroplorir Pied Butcherbird ** Anlhur novnescelondiw Richard's Pipit * A Tneniopygto gultato Zebra Finch A hephilo ncuticnudo Long-tailed Finch A Porphilo pcrronaln Masked Finch * A Emblemn picturn Painted Finch * A Erythrurn gouldiae Gouldian Finch Chcramoeco lcucortcmur White-backed Swallow ** ?Cinrlornmphur mthmi Rufous Songlark 33-38 1 'pursued, but not captured (Pettigrew ct ol. 1985). #listed as Cinrlorom cinnamomcum by Douglas (1967); name change due to taxonomic revision.

June 1999 Australian Zoologis1 31(1) 89 Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021

90 Australian Zoologist 31(1) June 1999 REFERENCES Higgins, P. J. and Davies, S. J. J. F. (eds), 1996. Handbook of Ausholian, Ncw Zeoland and Antontic Baird, R. El, 1991. The taphonomy of Late Quaternary Birds. Vol. 3. Term to Pigcmr. Oxford University cave localities yielding vertebrate remains in Press: Melbourne. Australia. Pp. 267-310 in Vcrtcbrntc Pnlurmtology of Aurholaria ed by Vickers-Rich, R. F. Baird, J. Kulzer, E., Nelson, J. E., McKean, J. L. and Maehres, F. Monaghan and T H. Rich. Thomas Nelson: P., 1984. Prey-catching behaviour and echolocation Melbourne. in the Australian Ghost Bat, Mncroderma gigas (Micmchimptera, Megadermatidae). Aut. . Boles, W. E., 1995. Preliminary analysis of the Passeri- 7: 37-50. former from Riversleigh, northwestern Queensland, Australia, with the description of a new species Marchant, S. and Higgins, P. J. (eds), 1993. Handbook of lyrebird. Cou"er Forsihunginstitut Smdenberg 181: of Austmlinn, New Zealand and Antantic Birdr. 163-70. Vol. 2. Roptorr to Lnpwings. Oxford University Press: Melbourne. Boles. W E., 1997. A kingfisher (Halcyonidae) from the Miocene of Riversleigh, northwestern Queensland, Newton, I., 1993. Predation and limitation of bird with comments on the evolution of kingfishers in numbers. Pp. 143-98 in Cumeni Ornithology Wl. 11 Australo-Papua. Mcm. Qld Mu. 41: 22S34. ed by D. M. Power. Plenum Press: New York.

Baler, W. E., 1998. A Budgerigar Mdopdtlo~urundulatur Pettigrew, J., Bakr, G. B., Baker-Gabb, D., Baverstock, Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/31/1/82/1474965/az_1999_009.pdf by guest on 24 September 2021 from the Pliocene of Riversleigh, northwestern G., Coler, R., Conole, L., Churchill, S., Fitzherbert, Queensland. Emu 98: 32-35. K., Guppy. A., Hall, L., Helman, P., Nelson, J., Priddel, D., Pulsford, I., Richards, G., Schulz, M. Douglas. A. M., 1967. The natural history of the and Tidemann, C. R., 1985. The Australian Ghost Ghost Bat, Mocrodcm gigas (Microchiraptera, Bat, Mocrodcrma gigar, at Pine Creek, Northern Megadermatidae), in Western Australia. WA Nal. Territory. Morrodemn 2: 8-19. 10: 125-38. Richards, G. C. and Hand, S., 1995. Ghost Bat Ford, H. A,, 1989. Ecology of Birds. An Australian Macroderma gigas. Pp. 446-47 in The Mammnlr of Pmpcctiue. Surrey Beatty & Sons Pty Ltd: Chipping Aurtralio ed by R. Strahan. Australian Museum and Norton, Sydney. Reed Books: Sydney. Guppy, A. and Coles, R. B., 1983. Feeding behaviour Schulz. M., 1986. Vertebrate prey of the Ghost Bat, of the Australian Ghost Bat, Mucro&mo gigas Mncrodcm gigas, at Pine Creek, Northern Territory. (Chiroptera: Megadermatidae) in captivity. Aud. Mamo&~ma 2: 59-62. Mammal. 6: 97-99. Sullivan, K. A. and Roper, J. J., 1996. Impacts of Hall, B. P. (ed), 1974. Birdr of the Harold Hall predation on passerine post-fledging success. Australian Expxpcditionr, 1962-70. British Museum Transactions of the 61" North American Wildl$e and (Natural History): London. Natural Resoume Conference, 77-85. Hand, S. J., 1985. New Miocene megadermatids Tidemann, C. R., Priddel, D. M., Nelson, J. E. (Chiroptera: Megadermatidae) from Australia with and Pettigrew, J. D., 1985. Foraging behaviour comments on megadermatid phylogenetics. Aust. of the Au3tralian Ghost Bat, Macrodrrma gigns Mamml. 8: 5-43. (Microchiroptera: Megadermatidae). Aust. j. Zool. 705-13. Hand, S. J., 1990. Australia's first Tertiary molossid 33: (Micmchiroptera: Molossidae): its phylogenetic and Toop, J., 1985. Habitat requirements, survival strategies biogeographic implications. Mcm. QU MM. 28: and ecology of the Ghost Bat Mawode- gigas 175-92. Dabsan, (Microchiroptera, Megadermatidae) in central coastal Queensland. Mncrodrrmn 1: 3741. Hand. S. 1.. 1995. First record of the renus Mecndcnna deoff;iy (Microchiroptera: ~e~a~rmatid;)from Vestjens, W J. M. and Hall, L. S., 1977. Stomach Australia. hlarovrrtdrato 24: 47-66. contents of forty-two species of bats from the Australasian region. 4: 25-35. Hand, S. J., 1996. New Miocene and Pliocene Awl. Wildl. Rer. megadermatids (Mammalia, Microchiroptera) from Australia, with comments on broader aspects of megadermatid evolution. Gcobior 49: 365-77. Hand, S. J., Dawsan, L. and Augee, M., 1988. Macvodennn koppo, a new Pliocene species of false vampire hat (Microchiroptera: Megadermatidae) from Wellington Caves, New South Wales. &c. Aurt. Mur. 40: 343-51.

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