Pteropus tonganusPacific flying fox

Ge­o­graphic Range

Ptero­pus ton­ganus is re­stricted to is­lands in the Pa­cific Ocean. It can be found dis­trib­uted widely from is­lands near New Guinea through the Cook Is­lands. (Grant, et al., June 1997)

Habi­tat

Ptero­pus ton­ganus lives in a va­ri­ety of habi­tats. These bats are typ­i­cally found in low­land or mon­tane na­tive forests. Bare, upper branches of very tall trees in ma­ture, pri­mary for­est are fa­vored as a roost­ing habi­tat. How­ever, roosts can lo­cated on cliff faces above the ocean or steep moun­tain­sides. These bats are also known to use coastal swamps, man­grove swamps, in­land swamp and marsh areas, as well as agri­cul­tural areas and human res­i­den­tial areas. (Brook, et al., June 2000; Miller and Wil­son, 1997)

Phys­i­cal De­scrip­tion

Ptero­pus ton­ganus is a black-backed bat with a con­trast­ing or­ange or yel­low­ish man­tle, re­sem­bling many other sym­patric species of bats. The total length of these bats av­er­ages 231 mm, and the mass av­er­ages 565 g. The fore­arm length, use­ful in help­ing dis­tin­guish P. ton­ganus from sym­patric bat species, is be­tween 120 and 160 mm, and the ears are al­ways less than 31 mm in length. Some re­searchers think that males grow to be larger than fe­males, but fur­ther in­for­ma­tion on this pos­si­ble sex­ual di­mor­phism is not pre­sent in the lit­er­a­ture. (Miller and Wil­son, 1997)

Sep­a­ra­tion of P. ton­ganus from a sim­i­lar species, P. samoen­sis, is made based on tooth pat­terns. Ptero­pus samoen­sis has a shorter, broader ros­trum than does P. ton­ganus. Ptero­pus samoen­sis is also much larger than P. ton­ganus. It is eas­ily dis­tin­guished from P. ton­ganus both by its larger size and by the ab­sence of the white fur patch on the back of its neck which char­ac­ter­izes P. ton­ganus. The pres­ence of a buffy man­tle on the neck and upper back of P. ton­ganus is the only di­ag­nos­tic char­ac­ter­is­tic that dis­tin­guishes it from P. samoen­sis in the field. (Ba­nack, Sep­tem­ber 1998; Cox, 1983; Wil­son and En­g­bring, 1992)

  • Sexual Dimorphism
  • male larger
  • Range mass
    191 to 1,099 g
    6.73 to oz
  • Average mass
    565 g
    19.91 oz
  • Range length
    151 to 262 mm
    5.94 to 10.31 in
  • Average length
    231 mm
    9.09 in

Re­pro­duc­tion

The mat­ing sys­tem of these an­i­mals is not doc­u­mented.

Adult fe­males gen­er­ally have only one young per year, with a ges­ta­tion of 140-192 days (4.6-6.3 months). The tim­ing of the breed­ing sea­son seems to vary some ge­o­graph­i­cally, and there are re­ports that these an­i­mals may breed year round in Amer­i­can Samoa.

Moth­ers carry their young in­fants with them, tucked be­neath a wing. Ob­served cop­u­la­tions often in­volve fe­males who are nurs­ing young, in­di­cat­ing that there may be a post-par­tum es­trus cycle.

Young begin to fly when they are ap­prox­i­mately three months old, and when they are only half to 3/4 of adult size. Weaned in­di­vid­u­als are often seen fly­ing and for­ag­ing with adults.

The young are usu­ally not weaned until they are 4-6 months old, and can re­main de­pen­dent on their moth­ers for a year. These an­i­mals do not reach sex­ual ma­tu­rity until they are 1.5- 2.0 years old. (Cox, 1983; Falan­ruw, 1998; Miller and Wil­son, 1997)

Al­though males and fe­males as­so­ci­ate with one an­other for most of the year, after mat­ing and be­com­ing preg­nant, fe­males are known to sep­a­rate them­selves from males to form "ma­ter­nity camps." When the young are weaned, the fe­males may re­join the males. (Miller and Wil­son, 1997)

  • Breeding interval
    These animals reproduce once per year.
  • Breeding season
    Breeding season apparently varies geographically, with some populations breeding in June or July, and others possibly breeding year round.
  • Range number of offspring
    1 to 2
  • Range gestation period
    140 to 192 days
  • Range weaning age
    4 to 6 months
  • Range age at sexual or reproductive maturity (female)
    1.5 to 2 years
  • Range age at sexual or reproductive maturity (male)
    1.5 to 2 years

The parental care of P. ton­ganus as not been well doc­u­mented. How­ever, it is known that fe­males pro­vide their off­spring with milk. The fe­male car­ries her off­spring when it is very small. Weaned off­spring often for­age with adults, which might be their par­ents. It is not known what role males of this species play in parental care. (Cox, 1983; Miller and Wil­son, 1997)

  • Parental Investment
  • altricial
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • post-independence association with parents

Lifes­pan/Longevity

Sur­vival rates of Ptero­pus in the wild are un­known, but the max­i­mum life span for a cap­tive an­i­mal is 31 years. (Pier­son and Rainey, 1992)

  • Range lifespan
    Status: captivity
    31 (high) years

Be­hav­ior

Ptero­pus ton­ganus is pri­mar­ily noc­tur­nal. This species is gre­gar­i­ous, and may be found in large groups while for­ag­ing, and dur­ing day­time, when the bats gather at large colo­nial roosts, often with sev­eral hun­dred bats in a sin­gle tree. Colonies are ex­tremely noisy and have a dis­tinct odor that makes them easy to de­tect. (Brook, et al., June 2000; Cox, 1983; Craig, et al., Oc­to­ber 1994)

When roost­ing, these bats hang up­side-down from their feet, wrap­ping their wings around them­selves as a shel­ter against in­clem­ment weather. The feet are gen­er­ally kept close to­gether. When mov­ing around in the roost, these bats are able crawlers, using their wings to drag them­selves along. (Miller and Wil­son, 1997)

Ptero­pus ton­ganus uti­lizes a broad array of plants for food. It takes pollen, nec­tar, and fruit from these trees. For­ag­ing typ­i­cally be­gins after dark, al­though some pop­u­la­tions have been known to for­age well be­fore sun­set. Such pop­u­la­tions may be under hunt­ing pres­sure, or food short­ages may pro­duce a need for day­time for­ag­ing. (Miller and Wil­son, 1997)

While for­ag­ing, these bats may travel a long way from their roost­ing sites. They do not al­ways eat their meals at the for­ag­ing site, though, and are known to carry large pieces of fruit away from their trees of ori­gin. The bats ma­nip­u­late the fruits with their hind legs, then smash the fruit against their palate to ex­tract the juices and pulp, which are swal­lowed, and eject the seeds. (Miller and Wil­son, 1997)

Ptero­pus ton­ganus are known to de­fend food re­sources. They may chase, bat at, and vo­cal­ize at in­trud­ing bats to warn them away. (Miller and Wil­son, 1997)

The so­cial struc­ture of this species is not well un­der­stood. Males and fe­males are some­times found in the same colonies, but fe­males sep­a­rate from the males to form "ma­ter­nity camps", only re­join­ing the males when the young are weaned. (Miller and Wil­son, 1997)

Home Range

Home range size for this species has not been re­ported.

Com­mu­ni­ca­tion and Per­cep­tion

These bats are known to have high pitched vo­cal­iza­tions. Al­though not specif­i­cally men­tioned in the lit­er­a­ture, be­cause of the way they roost so close to one an­other, these is un­doubt­edly some tac­tile com­mu­ni­ca­tion be­tween an­i­mals when roost­ing. The mother also has tac­tile com­mu­ni­ca­tion with her off­spring, whom she car­ries. There are re­ports of P. ton­ganus chas­ing one an­other in the con­text of de­fend­ing a food re­source, and bat­ting at one an­other, in­di­cat­ing fur­ther vi­sual and tac­tile com­mu­ni­ca­tion. (Cox, 1983; Miller and Wil­son, 1997)

Food Habits

P. ton­ganus is a gen­er­al­ist among fly­ing foxes, in that it con­sumes fruit, flow­ers, nec­tar, and pollen. Some­times, these an­i­mals have been known to eat leaves.

Fly­ing foxes swal­low mostly the juices ex­tracted after mas­ti­ca­tion of the fruit pulp and dis­card most of the fiber (in the form of an ejecta pel­let). These ejecta pel­lets are made as a bat feeds on fruit pulp, com­press­ing the fruit against the palate with the tongue. This ac­tion ef­fec­tively squeezes out the juices, which are swal­lowed, and cre­ates a pel­let of dry fruit pulp. Some of this pulp is swal­lowed, but the an­i­mals spit most of it out.

Most fruit con­sumed by fly­ing foxes have large seeds. These an­i­mals do not gen­er­ally con­sume fruits until they are fully ripe, as that is when the fruits are most juicy, and eas­i­est to smash up. They will con­sume un­ripe fruit in times of low avail­abil­ity of pre­ferred fruit re­sources.

The plant species most com­monly used for fruit in Samoa by P. samoen­sis and P. ton­ganus are Ar­to­car­pus al­tilus, Plan­chonella samoen­sis, and Syzy­gium ino­phyl­loides. P. ton­ganus is an ex­treme gen­er­al­ist and likely feeds upon more than 70 plant species. Be­cause it has so many sources for food, this species is well pre­pared to pe­riod en­vi­ron­men­tal dis­rup­tions, such as hur­ri­canes.

After a major dis­tur­bance, such as a hur­ri­cane, these an­i­mals may eat leaves.

P. ton­ganus is ex­tremely se­lec­tive in choos­ing fruit within a tree. An­i­mals will smell and oc­ca­sion­ally bite many fruits be­fore ei­ther eat­ing. They may eat the fruit in the tree, or may re­move one to eat in an­other lo­ca­tion.

P. ton­ganus can be an agri­cul­tural pest, for­ag­ing on bread­fruit, pa­payas, man­gos, ba­nanas and seasea fruits. Be­cause they do not dam­age cash crops, how­ever, they are thought mainly to be nui­sances, and not dam­ag­ing, by the Samoans. (Ba­nack, Sep­tem­ber 1998; Cox, 1983; Miller and Wil­son, 1997; Nel­son, et al., Oc­to­ber 2000)

  • Plant Foods
  • leaves
  • fruit
  • nectar
  • pollen
  • flowers

Pre­da­tion

Ptero­pus ton­ganus has few nat­ural preda­tors, partly be­cause of its in­su­lar lo­ca­tion. (Brautigam and Elmqvist, 1990)

Rap­tors and snakes are thought to be some of the major preda­tors of fly­ing foxes on is­lands. Pere­grine fal­cons, Falco pere­gri­nus, are known to sub­sist al­most ex­clu­sively on fly­ing foxes in New Cale­do­nia , the Loy­alty Is­lands, the Solomon Is­lands, and Van­u­atu. Barn owls also have been re­ported to take P. ton­ganus. (Miller and Wil­son, 1997)

Ecosys­tem Roles

Ptero­pus ton­ganus is an im­por­tant as pol­li­na­tor and seed dis­perser in Pa­cific is­land ecosys­tems. With­out them, it would be dif­fi­cult to main­tain com­mu­nity di­ver­sity, be­cause of the dis­tur­bances by storms. These an­i­mals are needed to af­fect the re­gen­er­a­tion of dom­i­nant for­est trees. They are also nec­es­sary to main­tain ge­netic flow be­tween is­lands. The loss of fly­ing foxes may af­fect plants that have co­e­volved with them. About 30 per cent of plant species on Samoa are to­tally de­pen­dent on fly­ing foxes for pol­li­na­tion or seed dis­per­sal. This is re­mark­ably high com­pared with forests in con­ti­nen­tal areas. Fly­ing foxes like P. ton­ganus are key­stone pol­li­na­tors and seed dis­persers, and dis­rup­tions of their pop­u­la­tions would re­sult in chain re­ac­tion of other ex­tinc­tions--of plants and other an­i­mals that are de­pen­dent upon them. (Ba­nack, Sep­tem­ber 1998; Brautigam and Elmqvist, 1990; Craig, et al., Oc­to­ber 1994; Miller and Wil­son, 1997)

Mu­tu­al­ist Species
  • Ceiba pen­tan­dra
  • Rhus tait­en­sis
  • Canan­gus idi­rata
  • Cocos nu­cifera
  • Ery­th­rina varia­gata
  • Musa sp.
  • Syzy­gium clu­si­ae­folium
  • Syzy­gium ino­phyl­loides
  • Syzy­gium malac­cense
  • Syzy­gium richii
  • Pan­danus tec­to­rius
  • Plan­chonella tor­ri­cel­len­sis
  • Dra­con­tomelon sp.
  • Mangifera in­dica
  • Cer­bera mang­has
  • Neisosperma op­posi­ti­folium
  • Car­ica pa­paya
  • Ter­mi­na­lia cat­apa
  • Diospy­ros samoen­sis
  • Elae­icar­pus raro­ton­gen­sis
  • Calo­phyl­lum ino­phyl­lum
  • Persea amer­i­cana
  • In­ocar­pus fag­ifer
  • Fa­graea bert­eroana
  • Ar­to­car­pus al­tilis
  • Ar­to­car­pus het­ero­phylla
  • Ficus co­piosa
  • Ficus pro­lixa
  • Ficus obli­qua
  • Myris­tica hy­par­gy­raea
  • Psid­ium gua­java
  • Cit­rus sinen­sis
  • Pom­me­tia pin­nata
  • Plan­chonella grayana
  • Plan­chonella samoen­sis
  • Lyciper­si­con es­cu­len­tum

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

These an­i­mals are hunted by local peo­ples for sub­sis­tence. Such hunt­ing is sus­tained year-round. They are also com­mer­cially hunted and over­har­vested on many Pa­cific Is­lands to sup­ply an ex­otic food mar­ket in Guam and for med­i­c­i­nal pu­poses. (Craig, et al., Oc­to­ber 1994)

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Be­cause of their feed­ing habits, these an­i­mals can po­ten­tially do dam­age to crops. (Miller and Wil­son, 1997)

  • Negative Impacts
  • crop pest

Con­ser­va­tion Sta­tus

Be­cause of com­mer­cial and sub­sis­tance hunt­ing, this species is some­times sub­ject to over­hunt­ing. Be­cause of their key role in local ecosys­tems, the po­ten­tial dam­age to the area from de­creases in pop­u­la­tion size are ex­treme.

Since 1989, P. ton­ganus has been listed in CITES Ap­pen­dix I. The Ap­pen­dix I list­ing pro­vides for a pro­hi­bi­tion on in­ter­na­tional trade in those species that are listed. How­ever, the species is not listed as en­dan­gered, threat­ened, or oth­er­wize at risk by the US Fish and Wildlife Ser­vice, or by the IUCN Redlist. (Brautigam and Elmqvist, 1990; Miller and Wil­son, 1997)

Other Com­ments

Ptero­pus ton­ganus plays an im­por­tant role in the eth­no­bi­ol­ogy of the Samoan peo­ple. The leg­ends of these peo­ple say that the Queen, Leu­togi Tupa’itea, had dis­pleased her hus­band, the king of Tonga. She was bar­ren, so he chased her into the crotch of a Cal­lo­phy­lum ino­phyl­lum tree. The vil­lagers promptly set a huge fire be­neath the tree. Just as the flames reached the Queen, a large flight of P. ton­ganus put out the flames. (Cox, 1983)

Con­trib­u­tors

Leslie Ava­los (au­thor), Hum­boldt State Uni­ver­sity, Brian Ar­bo­gast (ed­i­tor), Hum­boldt State Uni­ver­sity.

Nancy Shef­ferly (ed­i­tor), An­i­mal Di­ver­sity Web.

Glossary

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

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

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.

fertilization

union of egg and spermatozoan

folivore

an animal that mainly eats leaves.

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.

island endemic

animals that live only on an island or set of islands.

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).

keystone species

a species whose presence or absence strongly affects populations of other species in that area such that the extirpation of the keystone species in an area will result in the ultimate extirpation of many more species in that area (Example: sea otter).

marsh

marshes are wetland areas often dominated by grasses and reeds.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

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

nectarivore

an animal that mainly eats nectar from flowers

nocturnal

active during the night

oceanic islands

islands that are not part of continental shelf areas, they are not, and have never been, connected to a continental land mass, most typically these are volcanic islands.

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.

scrub forest

scrub forests develop in areas that experience dry seasons.

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

social

associates with others of its species; forms social groups.

suburban

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

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

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.

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.

year-round breeding

breeding takes place throughout the year

Ref­er­ences

Ba­nack, S. Sep­tem­ber 1998. Diet se­lec­tion and re­source use by fly­ing foxes (genus Ptero­pus). ECOL­OGY, 79(6): 1949-1967.

Brautigam, A., T. Elmqvist. 1990. Con­serv­ing Pa­cific is­land fly­ing foxes. ORYX, 24(2): 81-89.

Brook, A., C. Solek, A. Tu­alaulelei. June 2000. Roost­ing be­hav­ior of colo­nial and soli­tary fly­ing foxes in Amer­i­can Samoa (Chi­roptera: Pteropo­di­dae). Pa­cific Sci­ence, 54(4): 301-311.

Cox, P. 1983. Ob­ser­va­tions on the nat­ural his­tory of Samoan bats. MAM­MALIA, 47(4): 519-523.

Craig, P., T. Morell, P. Trail. Oc­to­ber 1994. The de­cline of fruit bats in Amer­i­can Samoa due to hur­ri­canes and over­hunt­ing. Bi­o­log­i­cal Con­ser­va­tion, 69: 261-266.

Falan­ruw, M. 1998. On the sta­tus, re­pro­duc­tive bi­ol­ogy and man­age­ment of fruit bats of Yap. Mi­cronesica, 21: 39-51.

Grant, G., P. Craig, P. Trail. June 1997. Cy­clone-in­duced shift in for­ag­ing be­hav­ior in fly­ing foxes in Amer­i­can Samoa. BIOTROP­ICA, 29(2): 224-228.

Miller, C., D. Wil­son. 1997. Ptero­pus ton­ganus. Mam­malian Species, 552: 1-6.

Nel­son, S., M. Miller, E. Heske, G. Fahey. Oc­to­ber 2000. Nu­tri­tional qual­ity of leaves and un­ripe fruit con­sumed as famine foods by the fly­ing foxes of Samoa. Pa­cific Sci­ence, 54(4): 301-311.

Pier­son, E., W. Rainey. 1992. The bi­ol­ogy of fly­ing foxes of the genus Ptero­pus: A re­view. US Fish & Wildlife Re­source Bi­o­log­ica Re­port, 90(23): 134-147.

Wil­son, D., J. En­g­bring. 1992. The fly­ing foxes Ptero­pus samoen­sis and Ptero­pus ton­ganus:sta­tus in Fiji and Samoa. US Fish and Wildlife Ser. Biol. Rep., 90(23): 36-38.