Pteropodidae

 Eastern tube-nosed bat Nyctimene robinsoni, photographed by Michael Pennay.

Belongs within: Chiroptera.
Contains: Nyctimenina, Pteropodina, Cynopterina, Dobsonia, Rousettus, Epomophorus.

The Cynopterini is a group of generally smaller fruit bats including the dog-faced fruit bats (Cynopterus) and the tube-nosed fruit bats (Nyctimene).

Fruit bats in Africa
Published 19 January 2016
A western Woermann’s fruit bat Megaloglossus azagnyi clings to the underside of a palm frond, copyright Jakob Fahr.

The diversity of bats often goes under-appreciated. With over 1200 known species around the world (with new ones continuing to be named on a regular basis), they significantly outnumber any of the other traditionally recognised orders of mammals except the rodents. The basal relationships among the bats have been subject to some disagreement in recent years but it is generally agreed that living bats can be divided between three main lineages, one of which is the Pteropodidae, the fruit bats of the Old World. Pteropodids differ in a number of ways from all other bats, the most notable of which being that they mostly do not use echolocation (some do, but not in the same way as other bats: whereas non-pteropodids use sounds produced by their vocal chords for sonar, echolocating pteropodids click with their tongues or clap their wings). Instead, pteropodids use their over-sized eyes to see their way in the dark. The most familiar pteropodids may be the large flying foxes of the genus Pteropus, but they are not the only members of the family.

Not all pteropodids are as large as the flying foxes, either. Megaloglossus is a genus of one or two species of fruit bat found in lowland rain forests in tropical Africa. They are the smallest of Africa’s pteropodids (forearm length is about four centimetres, which I’m guessing translates into a wingspan of about a foot?) and differ from other African pteropodids in their slender faces and long, protrusible tongue (the latter feature, of course, explaining the genus name). For a long time, Megaloglossus was classified with other long-tongued pteropodids in a subfamily Kiodotinae, of which it would have been the only African component. However, Bergmans (1997) reclassified it as part of a uniquely African tribe, the Myonycterini, on the basis of features such as partially webbed toes and a ventral collar of thick hair. The ‘kiodotines’ are now recognised as a polyphyletic assemblage that have evolved their protrusible tongues convergently, presumably to feed on flower nectar.

Woermann’s fruit bat Megaloglossus woermanni being handled on a stick by an interfering human, copyright Natalie Weber.

Until recently, only a single species of Megaloglossus was recognised: Woermann’s fruit bat M. woermanni, originally named for a specimen from Gabon. A separate subspecies, M. woermanni prigoginei, has been suggested for larger individuals in the eastern part of the larger country now called Congo but Bergmans (1997) noted that the type specimen of the species fell within the size range for ‘prigoginei‘. A recent molecular study of Myonycterini by Nesi et al. (2013) identified a genetic divide between specimens from Cameroon, Gabon and the Congos on one hand and Liberia and the Côte d’Ivoire on the other, and the authors proposed recognising the latter as a separate species Megaloglossus azagnyi for which they proposed the somewhat awkward vernacular name of ‘western Woermann’s fruit bat’. However, M. azagnyi is recognised solely on the basis of genetic distance; no morphological distinction has yet been identified between the populations. Nesi et al. (2013) claimed that specimens of M. azagnyi were generally smaller than M. woermanni, but their reported measurement ranges indicate a broad overlap between the two. Bergmans (1997), who examined a greater number of specimens than Nesi et al., suggested a broad cline of increasing size from the west to the east of Megaloglossus‘ range but no definite gap. Nesi et al.‘s division also leaves the status uncertain of populations in intervening regions such as Nigeria. The distribution of Megaloglossus is divided into two by the ‘Dahomey gap’, a region of dry coastal savannah in Benin, Togo and Ghana that splits the wetter rainforests on either side. However, as noted by Nesi et al. themselves, palaeontological records indicate that the Dahomey gap has not been consistenly present in the past, and it should not be assumed that it corresponds to the division between M. woermanni and M. azagnyi.

Systematics of Pteropodidae
<==Pteropodidae [Cephalotidae, Fructivorae, Frugivora, Harpyiae, Megachiroptera, Pteropi, Pteropidae, Pteropusideae]
    |  i. s.: Propotto Simpson 1967B78, S67
    |           `--*P. leakeyi Simpson 1967S67
    |--Archaeopteropus Meschinelli 1903SM93 [ArchaeopteropinaeKJ70]
    `--Pteropodinae (see below for synonymy)KJ70
         |  i. s.: Desmalopex Miller 1907M07
         |           `--*D. leucopterus Miller 1907M07
         |         Pterocyon Peters 1861 [incl. Leiponyx Jentink 1881]M07
         |           |--*P. stramineus (see below for synonymy)M07
         |           |--P. buettikoferi [=*Leiponyx buettikoferi]M07
         |           `--P. dupreanusM07
         |         Callinycteris Jentink 1889M07
         |           `--*C. rosenbergiiM07
         |         Odontonycteris Jentink 1902M07
         |           `--*O. meyeriM07
         |         Boneia Jentink 1879KJ70, M07
         |           |--*B. bidensM07
         |           `--B. menadensisM07
         |--+--EidolonFS15
         |  |    |--E. dupreanumIT07
         |  |    `--E. helvum (Kerr 1792)K92
         |  `--+--+--‘Pteropus’ insularisFS15
         |     |  `--‘Pteropus’ leucopterusFS15 [=P. (Spectrum) leucopterusM07]
         |     `--+--‘Pteropus’ subnigerFS15
         |        `--Pselaphon Gray 1870FS15, M07
         |             `--‘Pteropus’ pselaphon Layard 1829I92 [=P. (Spectrum) pselaphonM07; incl. *Pselaphon ursinusM07]
         `--+--+--NyctimeninaFS15
            |  `--Notopteris Gray 1859FS15, M07
            |       |--*N. macdonaldi Gray 1859M07, K92
            |       `--N. neocaledonica Trouessart 1908 [=N. macdonaldi neocaledonica]R14
            `--+--PteropodinaFS15
               `--+--CynopterinaFS15
                  `--+--+--+--‘Rousettus’ bidensFS15
                     |  |  `--+--Aproteles bulmeraeFS15
                     |  |     `--+--DobsoniaFS15
                     |  |        `--Harpyionycteris Thomas 1896FS15, M07 [Harpyionycterinae, Harpyionycterini]
                     |  |             |--*H. whiteheadi Thomas 1896M07, K92
                     |  |             `--H. celebensis Miller & Hollister 1921K92
                     |  `--+--Macroglossus Schinz 1824KJ70, M07 (see below for synonymy)
                     |     |    |--M. lagochilusBP87 [=Kiodotus lagochilusM07]
                     |     |    |    |--M. l. lagochilusM72
                     |     |    |    `--M. l. nanus [incl. M. lagochilus microtus, M. lagochilus pygmaeus]M72
                     |     |    |--M. minimus (Geoffroy 1810)MB12 [=Pteropus minimusM07, *Kiodotus minimusM07]
                     |     |    `--M. sobrinusIT07
                     |     `--Syconycteris Matschie 1899FS15, M07
                     |          |  i. s.: S. finschiM07 [=Kiodotus (Syconycteris) finschiM07, S. crassa finschiM72]
                     |          |         S. papuanaM07 (see below for synonymy)
                     |          |--S. hobbitFS15
                     |          `--+--*S. australis (Peters 1867)M07, FS15, K92 (see below for synonymy)
                     |             `--S. carolinaeFS15
                     `--+--+--Casinycteris argynnisFS15
                        |  `--Scotonycteris Matschie 1894FS15, M07
                        |       |--*S. zenkeriM07
                        |       |--S. bedfordiM07
                        |       `--S. ophiodonIT07
                        `--+--RousettusFS15
                           `--+--Eonycteris Dobson 1873FS15, M07
                              |    |  i. s.: E. rosenbergiBP87
                              |    |--*E. spelaea (Dobson 1871)M07, FS15, K92 [=Macroglossus spelaeusM07]
                              |    `--+--E. majorFS15
                              |       `--E. robusta Miller 1913FS15, K92
                              `--+--+--‘Rousettus’ lanosus Thomas 1906FS15, K92
                                 |  `--+--Megaloglossus Pagenstecher 1885FS15, M07 (see below for synonymy)
                                 |     |    `--M. woermanni [=*Trygenycteris woermanni]M07
                                 |     `--+--Lissonycteris angolensisFS15
                                 |        `--Myonycteris Matschie 1899FS15, M07
                                 |             |--M. relictaFS15
                                 |             `--+--*M. torquata (Dobson 1878)M07, FS15, K92 [=Rousettus torquatusM07]
                                 |                `--M. brachycephalaFS15 [=Xantharpyia brachycephalaM07]
                                 `--Epomophorini [Epomophorina]KJ70
                                      |--Plerotes anchietaeFS15
                                      `--+--Hypsignathus Allen 1861FS15, M07 (see below for synonymy)
                                         |    `--*H. monstrosus Allen 1861M07, K92 [incl. *Sphyrocephalus labrosusM07]
                                         `--+--Epomops Gray 1866FS15, M07
                                            |    |--E. dobsoniiFS15 [=Epomophorus (Epomops) dobsoniiM07]
                                            |    `--+--*E. franqueti (Tomes 1860)M07, FS15, K92 [=Epomophorus franquetiM07]
                                            |       `--E. buettikoferiFS15
                                            `--+--Nanonycteris Matschie 1899FS15, M07
                                               |    `--*N. veldkampii [=Epomophorus (*Nanonycteris) veldkampii]M07
                                               `--+--EpomophorusFS15
                                                  `--Micropteropus Matschie 1899KJ70, M07
                                                       |--*M. pusillus (Peters 1867)M07, K92 (see below for synonymy)
                                                       `--M. intermediusIT07

Hypsignathus Allen 1861FS15, M07 [incl. Sphyrocephalus Murray 1862M07, Zygaenocephalus (l. c.)M07]

Macroglossus Schinz 1824KJ70, M07 [=Macroglossa Lesson 1827M07; incl. Carponycteris Lydekker in Flower & Lydekker 1891M07, Kiodotus Blyth 1840M07, Rhynchocyon Gistel 1848 non Pteres 1847M07]

Megaloglossus Pagenstecher 1885FS15, M07 [incl. Trygenycteris Lydekker in Flower & Lydekker 1891M07]

*Micropteropus pusillus (Peters 1867)M07, K92 [=Epomophorus (Micropteropus) pusillusM07]

*Pterocyon stramineus [=Pteropus stramineus, Cynonycteris straminea, Xantharpyia (*Pterocyon) straminea; incl. Pterocyon paleaceus]M07

Pteropodinae [Carponycteriinae, Cephalotina, Cynopterini, Kiodotinae, Macroglossi, Notopterini, Pteropinae, Pteropini, Rousettina]KJ70

*Syconycteris australis (Peters 1867)M07, FS15, K92 [=Macroglossus minimus var. australisM07, Kiodotus (Syconycteris) australisM07; incl. K. (S.) crassusM07, Syconycteris crassaK92, S. naiasM72]

Syconycteris papuanaM07 [=Kiodotus (Syconycteris) papuanusM07, S. crassa papuanaM72; incl. S. crassa keyensisM72]

*Type species of generic name indicated

References

Bergmans, W. 1997. Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 5. The genera Lissonycteris Andersen, 1912, Myonycteris Matschie, 1899 and Megaloglossus Pagenstecher, 1885; general remarks and conclusions; annex: key to all species. Beaufortia 47 (2): 11–90.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[B78] Butler, P. M. 1978. Insectivora and Chiroptera. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 56–68. Harvard University Press: Cambridge (Massachusetts).

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[K92] Klompen, J. S. H. 1992. Phylogenetic relationships in the mite family Sarcoptidae (Acari: Astigmata). Miscellaneous Publications, Museum of Zoology, University of Michigan 180: i–vi, 1–154.

[KJ70] Koopman, K. F., & J. K. Jones Jr. 1970. Classification of bats. In: Slaughter, B. H., & D. W. Walton (eds) About Bats pp. 22–28. Southern Methodist University Press: Dallas.

[M72] McKean, J. L. 1972. Notes on some collections of bats (order Chiroptera) from Papua-New Guinea and Bougainville Island. CSIRO Division of Wildlife Research Technical Paper 26: 1–35.

[MB12] McKenzie, N. L., & R. D. Bullen. 2012. An acoustic survey of zoophagic bats on islands in the Kimberley, Western Australia, including data on the echolocation ecology, organisation and habitat relationships of regional communities. Records of the Western Australian Museum Supplement 81: 67–108.

[M07] Miller, G. S., Jr. 1907. The families and genera of bats. Smithsonian Institution, United States National Museum, Bulletin 57: i–xvii, 1–282, pls 1–14.

Nesi, N., B. Kadjo, X. Pourrut, E. Leroy, C. P. Shongo, C. Cruaud & A. Hassanin. 2013. Molecular systematics and phylogeography of the tribe Myonycterini (Mammalia, Pteropodidae) inferred from mitochondrial and nuclear markers. Molecular Phylogenetics and Evolution 66: 126–137.

[R14] Revilliod, P. 1914. Les mammifères de la Nouvell-Calédonie et des îles Loyalty. In: Sarasin, F., & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1 pt 4 pp. 341–365, pl. 10. C. W. Kreidels Verlag: Wiesbaden.

[S67] Simpson, G. G. 1967. The Tertiary lorisiform primates of Africa. Bulletin of the Museum of Comparative Zoology 136 (3): 39–62.

[SM93] Stucky, R. K., & M. C. McKenna. 1993. Mammalia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 739–771. Chapman & Hall: London.

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