Nyctimene robinsoniQueensland tube-nosed fruit bat

Ge­o­graphic Range

Queens­land tube-nosed fruit bats (Nyc­timene robin­soni) are found along the east coast of Queens­land, Aus­tralia rang­ing from north­ern New South Wales to the Cape York Penin­sula. Few ob­ser­va­tions have been made on the oc­cur­rence of Queens­land tube-nosed fruit bats south of the Queens­land bor­der but they are be­lieved to have a more scat­tered dis­tri­b­u­tion at the south­ern end of their range. Record­ings of the south­ern range of Queens­land tube-nosed fruit bats were taken south of Night­cap Na­tional Park at Snows Creek (a trib­u­tary of Coop­ers Creek found 26 km north north east of Lis­more, New South Wales) and around the for­est canopy at Boomerang Falls Flora Re­serve (about 5 km south south west of Snows Creek). Queens­land tube-nosed fruit bats are gen­er­ally dis­persed along the coast through­out their range; the far­thest in­land record­ing is also the most southerly record­ing and comes from the Cul­maran Creek val­ley in Rich­mond Range State For­est ap­prox­i­mately 84 km east of the coast. A spec­i­men housed by the Aus­tralian Mu­seum is re­ported to come from as far south as Wee Jasper, New South Wales, how­ever this is be­lieved to be at­trib­uted to a la­bel­ing error and this spec­i­men has been de­ter­mined to be is­land tube-nosed fruit bat. (Hall and Pet­ti­grew, 1995; Milledge, 1987; Riek, et al., 2010; Schulz, 1997)

Habi­tat

Al­though re­garded as a rain­for­est spe­cial­ist Queens­land tube-nosed fruit bats are found through­out com­plex no­to­phyll vine forests, Arau­car­ian no­to­phyll vine forests, mixed tall open forests, scle­ro­phyll veg­e­ta­tion, and in urban areas. Ac­counts of Queens­land tube-nosed fruit bats sug­gest that they pre­fer to roost amongst fo­liage in the rain­for­est sub-canopy layer where they find ef­fec­tive cam­ou­flage within the dried leaves. How­ever they have also been noted in ex­posed trees on for­est mar­gins. In­di­vid­u­als may roost alone from ap­prox­i­mately 4 to 6 m above the ground to higher up and out of sight in the canopy. In­di­vid­u­als show day-roost site fi­delity over short pe­ri­ods, mainly dur­ing ripe fruit abun­dances, within an area; but may change roost sites as a preda­tor avoid­ance strat­egy. Day-roost sites cho­sen by Queens­land tube-nosed fruit bats are vari­able and in­clude: pri­mary for­est sites, iso­lated trop­i­cal fig trees (Fis­cus ver­ie­gata), hind-dune man­groves, and sec­ond-growth forests near man­groves. (Richards, 1986; Schulz, 1997; Spencer and Flem­ing, 1989)

Phys­i­cal De­scrip­tion

The skull of the genus of tube-nosed fruit bats can be de­scribed as short and heavy with an­te­ri­orly deep ros­trum. Their lacrimal widths are a greater than dis­tance from orbit to nares. They have nar­row brain­case, slight ba­s­i­cra­nial flex­ion, and have an ave­o­lar line which pro­jects back­wards and passes through the condyle. Tube-nosed fruit bats have a non-tubu­lar oc­ciput and an­te­ri­orly fused pre­max­il­lar­ies. They have no nar­row­ing of the bony palate or only slight nar­row­ing be­hind max­il­lary toothrows. Tube-nosed fruit bats have par­al­lel upper ca­nines and the dis­tance be­tween pos­te­rior mo­lars is equal to width of in­terptery­goid fossa. They have an­gu­lar process of mandible that is greatly re­duced and long tubu­lar nos­trils. Their den­tal for­mula is in­cisors 1/0, ca­nines, 1/1, pre­mo­lars 3/3, and mo­lars 1/2 equal­ing 24. P2/ and P/2 are well de­vel­oped and equal in height to the cin­gu­lum of the ca­nines. The rest of the cheek­teeth are mo­lar­i­form all of which have high well-de­vel­oped an­te­rior cusps ex­cept for M/2, which is low crowned and about half the size of the other mo­lar­i­form teeth. Tongues of tube-nosed fruit bats have four cir­cum­val­late papil­lae ((An­ersen, 1912; Miller 1907) in Heaney and Pe­ter­son 1984). (Heaney and Pe­ter­son, 1984)

The den­ti­tion of the genus of tube-nosed fruit bats is unique be­cause they lack in­cisors on the lower jaw; the lower ca­nines func­tion in place of the miss­ing in­cisors. The mod­i­fied lower ca­nines nearly come into con­tact with each other and close against the 2 upper in­cisors when bit­ing. This odd den­ti­tion is at­trib­uted to a dis­tant an­ces­tor with re­duced bit­ing teeth, pos­si­bly due to a liq­uid based diet. In re­turn­ing to a diet of fruit a new bit­ing mech­a­nism evolved. (Hall, 1983)

Queens­land tube-nosed fruit bats have brown wings char­ac­ter­is­ti­cally speck­led with yel­low and lime-green spots. These spots pro­vide each in­di­vid­ual with their own “spot code” as no two bats have the same pat­tern. Their fur is grey to red-brown with a dark strip of fur cen­tered down the dor­sal side of their body. One of the most char­ac­ter­is­tic fea­tures of this fruit bat is their large bulging tubu­lar nos­trils (pro­trud­ing 5 to 6 mm from the face). This along with the char­ac­ter­is­tic yel­low to green spots on their wings, face, and ears, as well as their bulging eyes dis­tin­guishes them from all other Aus­tralian bats. (Hall and Pet­ti­grew, 1995; Hall, 1983; Nel­lett, 2007; Richards, 1986)

The wings of Queens­land tube-nosed fruit bats are short and broad com­pared to other mem­bers of the old world fruit bats fam­ily. Queens­land tube-nosed fruit bats gave a well-de­vel­oped tail and a claw on the index fin­ger (An­ersen, 1912; Miller 1907 as cited in Heaney and Pe­ter­son 1984). (Hall and Pet­ti­grew, 1995; Heaney and Pe­ter­son, 1984)

Queens­land tube-nosed fruit bats have a mea­sured basal meta­bolic rate of 54.7 cubic cm oxy­gen per h, and daily body tem­pa­tures rang­ing from 35 de­grees C to 37 de­grees C. Het­erothermy is rarely seen in the old world fruit bats fam­ily, how­ever Queens­land tube-nosed fruit bats have the abil­ity to rapidly drop their body te­per­a­ture, meto­bolic rate, and enter tor­por dur­ing day or night. Un­like other bat species which use shiv­er­ing or brown fat as their heat gen­er­at­ing mech­a­nisms, Queens­land tube-nosed fruit bats pro­duce heat much more rapidly via tachy­car­dia which is under ner­vous and hor­monal con­trol. (Hall and Pet­ti­grew, 1995; Riek, et al., 2010)

Queens­land tube-nosed fruit bats have an av­er­age head and body length of 100 to 110 mm as well as an av­er­age tail length of 20 to 25 mm. Com­monly used as a proxy for body con­di­tion by bat re­searchers the av­er­age fore­arm length and mass are 60 to 70 mm and 30 to 50 g re­spec­tively. Al­though lit­tle ev­i­dence of the sex­ual di­mor­phism in Queens­land tube-nosed fruit bats has been doc­u­mented, sex­ual di­mor­phism of the pelvic gir­dle has been sug­gested based on pal­pa­tions of adult bats. The pal­pa­tions of the pelvic gir­dle of Queens­land tube-nosed fruit bats in­di­cate that they may ex­hibit sim­i­lar sex­ual di­mor­phism to other species of Aus­tralian fly­ing foxes. Fe­males have open, V-shaped pelvic gir­dles and the males have closed, O-shaped pelvic gir­dles. Fe­males have also been noted as hav­ing a more lightly col­ored pelage than males while sub-adults have a more in­ter­me­di­ate col­oration. De­spite this the over­all color pat­terns and av­er­age mass for both sexes are the same. (Chap­man, et al., 1994; Hall, 1983; Heaney and Pe­ter­son, 1984; Riek, et al., 2010)

An analy­sis of the cel­lu­lar DNA con­tent of Queens­land tube-nosed fruit bats has re­vealed that they have fewer genes than any other mam­mal (small­est genome size). Their total genome size is ap­prox­i­mately half that of the human genome, which has im­pli­ca­tions about which parts of the genomes of hu­mans, or other mam­mals, are re­dun­dant. (Hall and Pet­ti­grew, 1995)

  • Sexual Dimorphism
  • sexes colored or patterned differently
  • sexes shaped differently
  • Range mass
    30 to 50 g
    1.06 to 1.76 oz
  • Range length
    100 to 110 mm
    3.94 to 4.33 in
  • Range wingspan
    60 to 70 mm
    2.36 to 2.76 in
  • Average basal metabolic rate
    54.7 cm3.O2/g/hr

Re­pro­duc­tion

Lit­tle is known about the mat­ing sys­tems of Queens­land tube-nosed fruit bats. (O'Brien, 1993)

Lit­tle is known about the re­pro­duc­tive bi­ol­ogy of Queens­land tube-nosed fruit bats; how­ever most species be­long­ing to the old world fruit bats fam­ily have long pe­ri­ods of mat­ing, long preg­nan­cies, and long lac­ta­tion pe­ri­ods, with the re­pro­duc­tive cycle last­ing around 12 months in many species. Like the ma­jor­ity of old world fruit bats species Queens­land tube-nosed fruit bats are sea­sonal breed­ers. (O'Brien, 1993)

Fe­male Queens­land tube-nosed fruit bats give birth to one pup be­tween Oc­to­ber and De­cem­ber. Close rel­a­tives Philip­pine tube-nosed fruit bats are found in pri­mary forests near water bod­ies and among upper canopy fo­liage which is sim­i­lar to the habi­tat re­quire­ments to Queens­land tube-nosed fruit bats. The sim­i­lar­i­ties be­tween these two species make Philip­pine tube-nosed fruit bats a de­cent proxy for the re­pro­duc­tion of Queens­land tube-nosed fruit bats. Hei­d­men (1987) found Philip­pine tube-nosed fruit bats also have a sin­gle young each year. Ges­ta­tion in Philip­pine tube-nosed fruit bats are around three and a half months long while lac­ta­tion lasts four months or longer. (Hall, 1983; Heaney and Pe­ter­son, 1984; Hei­de­man, 1987)

  • Breeding interval
    Queensland tube-nosed fruit bats breed once a year.
  • Breeding season
    Queensland tube-nosed fruit bats mate from July to September.
  • Average number of offspring
    1
  • Range gestation period
    3 to 3.5 months
  • Range weaning age
    4 (low) months

Queens­land tube-nosed fruit bats have long pe­ri­ods of lac­ta­tion and the sin­gle pup is car­ried by the mother until it grows rather large. No in­for­ma­tion is cur­rently avail­able about parental care in this genus. (Hall, 1983)

Lifes­pan/Longevity

Noth­ing is known about the longevity of Queens­land tube-nosed fruit bats. How­ever old world fruit bats live at least 30 years in wild and cap­tive set­tings. (Nowak, 1994)

Be­hav­ior

Queens­land tube-nosed fruit bats are be­lieved to roost ex­clu­sively soli­tar­ily. One study in cap­tiv­ity found they used ag­gres­sive be­hav­ior, such as vo­cal­ized threats and even phys­i­cal at­tacks, when held cap­tive with con­specifics, re­gard­less of sex. How­ever a dif­fer­ent study cap­tured an adult fe­male and sub-adult male in two dif­fer­ent nets and found these two bats to be more af­fil­ia­tive. They were ob­served in close con­tact, even em­brac­ing. Al­though Queens­land tube-nosed fruit bats roost ex­clu­sively in a soli­tary fash­ion, they have been found to feed in a more gre­gar­i­ous man­ner with mul­ti­ple in­di­vid­u­als feed­ing in a sin­gle tree. These con­flict­ing ob­ser­va­tions on the so­cial be­hav­ior of Queens­land tube-nosed fruit bats may be due to the costs as­so­ci­ated with clus­ter roost­ing, which would dampen the ef­fi­cacy of their cam­ou­flage. Such costs would not be im­posed while for­ag­ing in the dark and the ripeness of fruit on in­di­vid­ual trees would also re­sult in clus­ters of bats feed­ing on the same tree. Queens­land tube-nosed fruit bats have also been ob­served feed­ing si­mul­ta­ne­ously with spec­ta­cled fly­ing fox in the same tree. (Hall and Pet­ti­grew, 1995; Milledge, 1987; Richards, 1986; Spencer and Flem­ing, 1989)

The short broad wings of Queens­land tube-nosed fruit bats pro­vide them with a low as­pect ratio (wing length to width) which al­lows for great ma­neu­ver­abil­ity. This low as­pect ratio al­lows them to hover for sev­eral sec­onds and change the di­rec­tion they are fac­ing while hov­er­ing. This be­hav­ior is rare for bats and is usu­ally only seen in small light-weight glean­ers. (Hall and Pet­ti­grew, 1995)

Ob­ser­va­tions made of south­ern pop­u­la­tions of Queens­land tube-nosed fruit bats doc­u­mented be­hav­iors not re­ported in north­ern pop­u­la­tions (which have been more thor­oughly in­ves­ti­gated). These be­hav­iors in­clude the re­quire­ment to drink after pro­longed hot and dry weather, as well as a lack of ac­tiv­ity dur­ing the cooler months of the year. Also, it in­cludes the move­ment of in­di­vid­u­als into ad­ja­cent open forests and moist tall open forests for oc­ca­sional for­ag­ing and drink­ing. (Schulz, 1997)

  • Average territory size
    200 m^2

Home Range

Queens­land tube-nosed fruit bats roost soli­tar­ily in the sub-canopy of trees dur­ing the day and feed on nearby fruit­ing trees within 200 m of their day roost at night. Al­though gen­er­ally found for­ag­ing within 200 m of their day roost sites they have been cap­tured from 63 m to 1012 m from their roost­ing sites using mist nets. Queens­land tube-nosed fruit bats have been found to change their roost­ing and feed­ing lo­ca­tions in an op­por­tunis­tic man­ner as they will move into new areas when food avail­abil­ity changes. Al­though they do change their roost­ing or feed­ing sites, they will also re­main within a small area for pro­longed pe­ri­ods of time dur­ing local food abun­dances. (Spencer and Flem­ing, 1989)

Com­mu­ni­ca­tion and Per­cep­tion

The tubu­lar shaped nos­trils were once be­lieved to func­tion as snorkels to aid Queens­land tube-nosed fruit bats in feed­ing on messy fruit meals. How­ever ob­ser­va­tions made of Queens­land tube-nosed fruit bats while feed­ing have re­vealed that they are not messy eaters, in­di­cat­ing that the tubu­lar nos­trils must serve some other pur­pose. Fur­ther in­ves­ti­ga­tion into the tubu­lar nos­trils has re­vealed that the nos­trils can open and close and move in­de­pen­dently of each other. This open­ing and clos­ing of the nos­trils oc­curs in re­sponse to au­di­tory, vi­sual, and ol­fac­tory cues and al­lows Queens­land tube-nosed fruit bats to scan their sur­rounds via ol­fac­tion. Like stereo vi­sion from two eyes or stereo hear­ing from two ears the in­de­pen­dent func­tion­ing of the two nos­trils al­lows for stereo ol­fac­tion, pro­vid­ing in­de­pen­dent mea­sures of aroma con­cen­tra­tions. The use of mod­i­fied stereo ol­fac­tion sys­tem al­lows them to lo­cate and fol­low odor plumes in a three di­men­sional realm. This stereo ol­fac­tion sys­tem aides in find­ing pre­ferred fig fruits through­out the Aus­tralian rain­forests. (Hall and Pet­ti­grew, 1995; Hall, 1983; Richards, 1986; Schwab and Pet­ti­grew, 2005)

Queens­land tube-nosed fruit bats com­mu­ni­cate via char­ac­ter­is­tic whistling calls which also serve as a re­li­able in­di­ca­tion of their pres­ence within an area. (Hall, 1983)

Food Habits

Queens­land tube-nosed fruit bats are spe­cial­ist fru­gi­vores feed­ing mainly on 3 species of fig (Ficus co­piosa; Ficus no­dosa; Ficus var­ie­gata). They are also known to feed on lil­lyp­illy (Syzigium corni­flo­rum), the ex­otics guava and sour­sop (Psid­ium gua­java; An­nona muri­cata) around or­chards and ona pi­o­neer species (Fis­cus race­mos) around aban­doned pas­tures. The ma­jor­ity of in­di­vid­u­als found feed­ing on sour­sop are fe­male which has been at­trib­uted to the dif­fer­ent nu­tri­tional re­quire­ments of preg­nant and lac­tat­ing fe­males, par­tic­u­larly as sour­sop has a much higher fat and pro­tein con­tent than the na­tive figs. For­ag­ing be­hav­ior is re­stricted to the un­der­story where cau­li­flor­ous (trunk-fruit­ing) trees are vis­ited. Queens­land tube-nosed fruit bats gen­er­ally for­age on trees within close prox­im­ity to their day roosts. Fruit may be car­ried away to be eaten or con­sumed on site at the tree where the fruit was grow­ing. In­di­vid­u­als can carry fruit weigh­ing over half of their own body weight. In one in­stance a 54 g bat was recorded car­ry­ing a 30 g fig. Mul­ti­ple in­di­vid­u­als will feed si­mul­ta­ne­ously on the same tree and have even been noted feed­ing along with spec­ta­cled fly­ing foxes. (Hall and Pet­ti­grew, 1995; Richards, 1986; Spencer and Flem­ing, 1989)

  • Plant Foods
  • fruit
  • nectar
  • pollen
  • flowers

Pre­da­tion

By wrap­ping their wings around their body, the col­ored spots of their wings, face, and ears make Queens­land tube-nosed fruit bats cryp­tic when roost­ing dur­ing the day amongst dense sun-mot­tled fo­liage. The color pat­tern of them is well adapted for cam­ou­flage within the canopy as their dark dor­sal stripe also re­sem­bles the cen­ter rib and stem of a dead leaf. Their soli­tary roost­ing be­hav­ior may allow for en­hanced ef­fi­cacy of their cryp­tic col­oration, as larger groups would make this color pat­tern ap­pear more con­spic­u­ous. Queens­land tube-nosed fruit bats also limit their ac­tiv­ity dur­ing full moon phases; this may be at­trib­ut­able to preda­tor avoid­ance for vi­sual noc­tur­nal preda­tors such as owls. (Hall, 1983; Richards, 1986; Riek, et al., 2010)

  • Anti-predator Adaptations
  • cryptic

Ecosys­tem Roles

Queens­land tube-nosed fruit bats are im­por­tant seed dis­persers and pol­li­na­tors for the Aus­tralian ecosys­tem. They are the only small, un­der­story, seed-dis­pers­ing bat in Aus­tralian trop­i­cal forests. When com­pared to the trop­i­cal forests of south­east Asia and cen­tral Amer­ica, which have many species of small, un­der­story, fruit-eat­ing bats that share the task of dis­pers­ing seeds, the im­por­tance of the role played by Queens­land tube-nosed fruit bats in Aus­tralian rain­forests is quite ap­par­ent. (Hall and Pet­ti­grew, 1995; Nel­lett, 2007)

Queens­land tube-nosed fruit bats have been found with par­a­sitic mites (Meris­taspis spp.), which are com­mon wing mites of old World fruit bats. (Dom­row, 1967)

  • Ecosystem Impact
  • disperses seeds
  • pollinates
Com­men­sal/Par­a­sitic Species
  • par­a­sitic mites (Meris­taspis kole­nati)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Queens­land tube-nosed fruit bats aid in for­est suc­ces­sion as well as dis­per­sal and prop­a­ga­tion of figs, pro­vid­ing en­hanced eco­nomic value to old world trop­i­cal forests. (Mus­carella and Flem­ing, 2007)

  • Positive Impacts
  • food
  • produces fertilizer
  • pollinates crops

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

In­creas­ing num­bers of ex­otic fruit or­chards in re­gions where Queens­land tube-nosed fruit bats are pre­sent have re­sulted in an in­crease of re­ports of these bats dam­ag­ing or­chard fruits such as star­fruit (Aver­rhoa muri­cata; Aver­rhoa caram­bola). (Spencer and Flem­ing, 1989)

  • Negative Impacts
  • crop pest

Con­ser­va­tion Sta­tus

Queens­land tube-nosed fruit bats have been noted as being vul­ner­a­ble (NSW TSC act) due to human de­vel­op­ment. Much of this con­cern is re­lated to en­tan­gle­ment in barbed wire fences. In­creases in the num­ber of or­chards in Aus­tralia has led to in­creased con­flict with the farm­ing com­mu­nity. Farm­ers have been known to hang nets around or­chards to keep the bats away from fruit crops. Bats that be­come en­tan­gled in the nets are left to starve. There­fore un­der­stand­ing the bi­ol­ogy of Queens­land tube-nosed fruit bats is par­tic­u­larly im­por­tant for man­age­ment strate­gies around fruit or­chards in order to min­i­mize the dam­age to the fruit as well as the bats. (Booth, 2007; Spencer and Flem­ing, 1989)

Con­trib­u­tors

Steve Smith (au­thor), Uni­ver­sity of Man­i­toba, Jane Wa­ter­man (ed­i­tor), Uni­ver­sity of Man­i­toba, Laura Podzikowski (ed­i­tor), Spe­cial Pro­jects.

Glossary

Australian

Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

coastal

the nearshore aquatic habitats near a coast, or shoreline.

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

Ref­er­ences

Booth, C. 2007. Barbed Wire Ac­tion Plan. Queens­land Con­ser­va­tion, Bris­bane, 2: 1-17.

Chap­man, A., L. Hall, M. Ben­nett. 1994. Sex­ual Di­mor­phism in the Pelvic Gir­dle of Aus­tralian Fly­ing Foxes. Aus­tralian Jour­nal of Zo­ol­ogy, 42: 261-265.

Dom­row, R. 1967. Mite par­a­sites of small mam­mals from scrub ty­phus foci in Aus­tralia. Aus­tralian Jour­nal of Zo­ol­ogy, 15: 759-798.

Hall, L. 1983. Queens­land Tube-nosed Bat Nyc­timene robin­soni.. Pp. 286-287 in Com­plete book of Aus­tralian mam­mals.. North Ryde, NSW: Corn­stalk Pub..

Hall, L., J. Pet­ti­grew. 1995. The bat with the stereo nose. Aus­tralian Nat­ural His­tory, 24: 26-28.

Heaney, L., R. Pe­ter­son. 1984. A new species of tube-nosed fruit bat (Nyc­timene) from Ne­gros Is­land, Philip­pines (Mam­malia: Pteropo­di­dae). Oc­ca­sional pa­pers of the Mu­seum of Zo­ol­ogy, Uni­ver­sity of Michi­gan, 708: 1-16.

Hei­de­man, P. 1987. The re­pro­duc­tive ecol­ogy of a com­mu­nity of Philip­pine fruit bats (Pteropo­di­dae, Megachi­roptera). Doc­toral dis­ser­ta­tion, Uni­ver­sity of Michi­gan, 1: 205.

Milledge, D. 1987. Notes on the oc­cur­rence of the Queens­land Tube-nosed Bat Nyc­timene robin­soni in north-east­ern New South Wales.. Macro­derma, 3: 28-30.

Mus­carella, R., T. Flem­ing. 2007. The role of fru­giv­o­rous bats in trop­i­cal for­est suc­ces­sion.. Bi­o­log­i­cal Re­views, 82: 573-590.

Nel­lett, K. 2007. An­a­lyz­ing Site Loy­alty in Nyc­timene robin­soni at the End of the Dry Sea­son. ISP Col­lec­tion, 144: 1-31.

Nowak, R. 1994. Walker's Bats of the World. Bal­ti­more, MD: The Johns Hop­kins Uni­ver­sity Press.

O'Brien, G. 1993. Sea­sonal re­pro­duc­tion in fly­ing foxes, re­viewed in the con­text of other trop­i­cal mam­mals.. Re­pro­duc­tion, fer­til­ity and de­vel­op­ment, 5: 499-521.

Richards, G. 1986. Notes on the nat­ural his­tory of the Queens­land tube-nosed bat, Nyc­timene robin­soni.. Macro­derma, 2: 64-67.

Riek, A., G. Ko­rt­ner, F. Geiser. 2010. Ther­mo­bi­ol­ogy, en­er­get­ics and ac­tiv­ity pat­terns of the East­ern tube-nosed bat (Nyc­timene robin­soni) in the Aus­tralian trop­ics: ef­fect of tem­per­a­ture and lunar cycle.. Jour­nal of Ex­per­i­men­tal Bi­ol­ogy, 213: 25-57.

Schulz, M. 1997. Notes on the east­ern tube-nosed bat, Nyc­timene robin­soni, from the Rich­mond Range, north­east­ern New South Wales.. Aus­tralian Mam­mal­ogy, 20: 127-130.

Schwab, I., J. Pet­ti­grew. 2005. A choroidal sleight of hand.. British Jour­nal of Oph­thal­mol­ogy, 89: 1398.

Spencer, H., T. Flem­ing. 1989. Roost­ing and For­ag­ing Be­hav­ior of the Queens­land Tube-Nosed Bat, Nyc­timene-Robin­soni (Pteropo­di­dae)-Pre­lim­i­nary Ra­dio-Track­ing Ob­ser­va­tions.. Wildlife Re­search, 16: 413-420.