Field of Science

Showing posts with label Variamana. Show all posts
Showing posts with label Variamana. Show all posts

Gazelles and their Kin

Female steenbuck Raphicerus campestris, copyright Yathin S. Krishnappa.


Ever since biblical times, gazelles have been a byword for a kind of watchful elegance, always on guard against unwanted advances. It is not difficult to see how such an analogy arose: on their native savannah, gazelles are indeed always on the alert, wary of the threat of predators and quick to respond to alarm. It is a habit that has served them for millions of years.

The Antilopini are an assemblage of about thirty species of mostly smaller antelope found in Africa and Asia*. The smallest are the dikdiks of the genus Madoqua which may be only a foot or so in height and weight just a few kilos; the tallest, the dibatag Ammodorcas clarkei, stands about 90 cm at the shoulder and weighs about 30 kilograms. They are mostly associated with arid or semi-arid habitats: savannahs, deserts, steppes and the like. Some species form sizeable herds; others live solitary lives.

*Before I go too much further, I should note that J. K. Revell over at his site Synapsida has written a number of posts about bovids (antelopes, cattle, etc.) over the the past few years that I heartily recommend. To the best of my knowledge, he hasn't gotten to antilopins yet, so I should be safe on that front.

Female and male oribi Ourebia ourebi, copyright Bill Higham.


Modern researchers largely agree on dividing the Antilopini between four major lineages, recognised as subtribes. One contains a single species, the oribi Ourebia ourebi, a smaller species with short, straight horns found in eastern sub-Saharan Africa. The Raphicerina, including the dikdiks Madoqua, the steenbucks and grysbucks Raphicerus and the beira Dorcatragus megalotis, are similar small, short-horned species. The Raphicerina and oribi are solitary species with individuals maintaining exclusive territories (at least between members of the same sex). They advertise their territories through the use of defecation sites together with the marking of vegetation using scent glands in front of the eye. The Raphicerina are exclusively browsers, concentrating on higher-quality food sources; in contrast, the oribi is a grazer and consequently must occupy a larger territory than the other species. Females of Raphicerina and oribi are hornless; in most other Antilopini (with some exceptions noted below), horns are present in both sexes though the females' horns are shorter and more slender.

Przewalski's gazelles Procapra przewalskii, copyright Yilun Qiao.


The majority of the remaining Antilopini live in herds though males of most species will claim temporary territories during the breeding season as they attempt to gather harems of females. The central Asian gazelles of the genus Procapra are placed in their own subtribe; these are three pale, medium-sized species found on steppes and high-altitude grasslands between the Himalayan plateau and Mongolia. They have rear-swept horns that make them look a bit like a gazelle that is trying to pass itself as a goat. Procapra gazelles may not be immediately related within the Antilopini to the true gazelles in the largest of the four subtribes, the Antilopina. Until recently, most authors would have treated the great majority of the Antilopina species in the genus Gazella; however, questions about the monophyly of this genus in the broad sense have lead to the recognition of three separate genera of gazelles: Gazella sensu stricto, Nanger and Eudorcas. The Nanger species, which include the dama gazelle N. dama and Grant's gazelle N. granti, are relatively large gazelles with a conspicuous white rump that is absent in the other two genera. The genus Eudorcas includes perhaps the most familiar gazelle species, Thomson's gazelle E. thomsoni of Kenya and Tanzania, which forms much larger herds than other gazelle species.

Mhorr gazelles Nanger dama mhorr at Tierpark Hellabrunn in München, copyright Rufus46.


The remaining living Antilopina species are all placed in their own separate genera. The springbuck Antidorcas marsupialis of southern Africa also forms large herds that used to number in the tens of thousands before hunting and habitat loss reduced their population. Springbucks are best known, of course, for their habit of 'pronking', a mode of bounding with all four legs held stiff and landing simultaneously, most often seen when the animal is alarmed or at play. Pronking is not unique to springbucks (other gazelles do it too) but it is made particularly noticeable in this species by a crest of white hairs towards the rear of the back that is erected at the same time.

Springbuck Antidorcas marsupialis engaged in some pronking, copyright Hans Stieglitz.


In other species of Antilopina, only the males have horns. The gerenuk Litocranius walleri and dibatag Ammodorcas clarkei are two slender species found in eastern Africa that differ from other Antilopina in being browsers rather than grazers and maintaining permanent exclusive territories. Both these species habitually feed while standing erect on the hind legs, allowing them to browse at higher levels than they could otherwise; they are even able to walk about to a certain extent in this pose, albeit perhaps not in a manner that could be called graceful. Outside of Africa, the blackbuck Antilope cervicapra is found in grasslands and woodlands of the Indian subcontinent (there is also supposed to have been a small introduced population of them near Geraldton here in Western Australia, though it may have since been eradicated). Males of this species have long, spirally twisted horns; mature males are also the only 'blackbucks' that are actually black (at least dorsally) whereas females and young males are light brown.

Pair of juvenile dibatags Ammodorcas clarkei at a rescue centre, copyright F. Wilhelmi.


Perhaps the most distinctive member of the Antilopina, however, is the saiga Saiga tatarica. This is the only species that is known to never be territorial, forming large herds in its native habitat of the central Asian steppes (technically, the social habits of the little-studied dibatag are largely unknown but it would not be unreasonable to presume that they are similar to those of the gerenuk). It is more robust than other Antilopina species; indeed, there was long uncertainty about whether saiga are more closely related to gazelle or goats. The nostrils of saiga are inflated to a hanging proboscis that is usually presumed to function as protection for the respiratory tissues from the dust of their near-desert habitat. However, there may also be a display function involved; during the mating season, the proboscis of males becomes engorged while scent glands in front of the eyes produce pungent secretions (so maybe the function of the proboscis is actually to somehow protect the saiga from its own stench). Unfortunately, the saiga (among other Antilopini species) is currently regarded as critically endangered, with only a fragment of its historical population surviving. There was a time when the saiga was thought to be something of a conservation success story: after being almost wiped out in the early 1900s, populations built up to about two million by the 1950s. But in the last few decades, the combined effects of factors such as habitat loss, disease and the demand for their horns from everyone's favourite country to turn the extermination of endangered species into a pointless investment bubble have caused numbers to crash back down to an estimated 50,000 or so (as relayed by Wikipedia).

Pair of saiga Saiga tatarica, copyright N. Singh.


Fossil species have been assigned to the genus Gazella from as far back as the Miocene though there may be grounds for debating how many of them are true Gazella. For instance, Bärmann (2014) commented on preliminary results of a phylogenetic analysis including the Pakistani Miocene species G. lydekkeri (from the well-studied Siwalik deposits) that suggested that it might be placed outside the Antilopina crown group. Other fossils of Antilopini inform us that the modern blackbuck is the sole survivor of a lineage of spiral-horned antelopes that was previously more widespread in Eurasia. The saiga was more widespread in the past as well, with either the modern or a closely related species known during the Pleistocene from more northerly parts of Siberia (at which point, presumably, there may have been saiga in the taiga) and even in northernmost North America. If they do disappear completely, it will be a sad end to a long history.

REFERENCES

Bärmann, E. V. 2014. The evolution of body size, horn shape and social behaviour in crown Antilopini—an ancestral character state analysis. Zitteliana B 32: 185–196.

Macdonald, D. (ed.) 1984. All the World's Animals: Hoofed Mammals. Torstar Books: New York.

Fruit Bats in Africa

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.

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.

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.