Voley, Voley, Voley

Over a third of all living mammal species are rodents. In cooler regions of the Northern Hemisphere, the rodent fauna is often dominated by the Microtinae, the group of mouse-like rodents including voles and lemmings. And in North America, the most widespread of all microtine species is the eastern meadow vole Microtus pennsylvanicus.

Eastern meadow vole Microtus pennsylvanicus, copyright Gilles Gonthier.

The eastern meadow vole is found over most of Canada and a large part of the northern and eastern United States, with the subspecies M. p. chihuahuensis known from Chihuahua in northern Mexico. This species is about the size of a small rat, being from 14 to 20 cm in length with about three to six centimentres of that length being tail (Reich 1981). They are generally yellowish-brown in colour with black tips on the hairs though individuals vary significantly in brightness and shade. Western populations are supposed to be lighter in coloration than eastern, and southern individuals tend to be larger than northern. As an indication of this species' variability, Reich (1981) recognised 28 recognised subspecies.

Eastern meadow voles are primarily inhabitants of grasslands, with a preference for damper habitats, though they may also be found in woodlands. They mostly live in burrows underground, emerging to the surface to forage for food. Eastern meadow voles are generalist feeders, browsing on most available forms of low vegetation: grasses, sedges and herbs. When populations reach their peak, they may cause significant damage to woody plants by ringbarking their trunks. Individuals may seemingly be active at just about any time of day.

Eastern meadow vole in a state of danger, copyright David Allen.

Like other small rodents, meadow voles are short-lived animals with estimates of average lifespan ranging from just two or three months to ten to fourteen months (Reich 1981). Studies of movement patterns indicate that mature females generally maintain distinct, non-overlapping ranges whereas males range further and with less concern for others (Madison 1980). Mating behaviour appears generally promiscuous: males will range over the territories of multiple females and litters with mixed paternity are not uncommon (Boonstra et al. 1993). Paternal behaviour has been observed among eastern meadow voles in laboratory populations but all indications are that wild males do not remain with females after mating. Males often bear wounds indicative of intra-species conflict. These may be the result of males fighting over access to females but Madison (1980) suggested a potential alternative. Less dominant males might be more likely to attempt to approach females earlier or later in their oestrus cycle as the females are more likely to be guarded by dominant males when at their peak. While avoiding attacks from their dominant brethren, these minor males might find themselves violently rebuffed by a female who is just not yet in the mood.

After mating, gestation lasts for about three weeks, usually resulting in a litter of four to six babies. Weaning then takes place after about two weeks. Females forage far less while lactating than at other times. It might seem counter-intuitive for a female to reduce feeding when her energy demands are presumably at their peak but again Madison (1980) suggests an explanation: perhaps her energy needs are such that she simply lacks the capacity for extensive wandering. Young may potentially remain with their mother for some time after weaning but eventually they will be forced out of the parental burrow, leaving to face the wide world on their own. And when you're the size of a vole, that's a very wide world indeed.

REFERENCES

Boonstra, R., X. Xia & L. Pavone. 1993. Mating system of the meadow vole, Microtus pennsylvanicus. Behavioral Ecology 4: 83–89.

Madison, D. M. 1980. Space use and social structure in meadow voles, Microtus pennsylvanicus. Behavioral Ecology and Sociobiology 7: 65–71.

Reich, L. M. 1981. Microtus pennsylvanicus. Mammalian Species 159: 1–8.

The Long-eared Bats of Australasia

When most people think of Australian mammals, they imagine the fauna as dominated by marsupials and monotremes, representatives of lineages long isolated from those found elsewhere. But Australia is also home to a remarkable diversity of native placentals. Immigrating from the north as Australia drifted closer to Asia, the rodents and bats underwent their own radiations on the Australian continent and its neighbouring islands. Among these distinctly Australasian assemblages of placentals are the long-eared bats or big-eared bats of the tribe Nyctophilini.

Lesser long-eared bat Nyctophilus geoffroyi, copyright Michael Pennay.

The long-eared bats comprise fifteen or so species found over a range between eastern Indonesia and Australia with outlying species in New Caledonia and Fiji. They are members of the Vespertilionidae, the most diverse recognised family of bats, and share with most other vespertilionids a fairly generalised appearance with dull coloration. They differ from other vespertilionids in having a relatively short muzzle (with a correspondingly reduced number of teeth) with a small nose-leaf at its end (Miller 1907). They also (as the vernacular name indicates) have particularly large ears, as long as or longer than the rest of the head, that are commonly connected medially by a distinct membrane. At rest, the ears may be folded like a concertina along the hind margin to protect them from damage (Hall & Woodside 1989). Historically, the long-eared bats were treated as their own subfamily within the Vespertilionidae that also included a similar North American genus Antrozous. However, the nyctophilins are now regarded as a derived tribe within the larger subfamily Vespertilioninae (albeit one whose exact relationships remain uncertain) and similarities between Nyctophilini and Antrozous are thought to be convergent rather than reflecing a close relationship. The majority of nyctophilins are placed in a single genus Nyctophilus with the exception of the New Guinea big-eared bat Pharotis imogene. This species differs from Nyctophilus in lacking hair at the end of the muzzle.

Nyctophilins are found in a range of habitats but seem to prefer dry woodlands. Vespertilionids as a whole are differentiated from other bats by modifications of the fore arms including a highly developed double joint between scapula and humerus and reduction of the ulna. As a result, they may be less powerful fliers than other bats but they would be more agile. This trend would be particularly pronounced in nyctophilins which have relatively short wings compared to other vespertilionids (Hall & Woodside 1989). The development of a nose-leaf in nyctophilins is associated with their use of signals emitted at a constant frequency through the nose for echolocation whereas other vespertilionids use signals of varying frequency emitted through the mouth. As well as catching insect prey in flight, long-eared bats are able to recognise prey at rest and so glean insects off vegetation or on the ground. This gleaning habit is presumably also associated with long-eared bats having relatively larger eyes than other vespertilionids.

Gould's long-eared bat Nyctophilus gouldi with ears partially reclined, copyright Department of Environment and Primary Industries, Victoria.

Caves in Australia are mostly not very extensive so the formation of colonies by Australian vespertilionids is constrained by the availability of suitable roosting sites such as hollows in trees or crevices in rocks. At least some long-eared bats may be solitary (Hall & Woodside 1989). Their distribution in Australia (as with pretty much all Australian animals) is also largely contingent on the availability of water. Mating happens in autumn but gestation is generally delayed, whether by delaying fertilisation or development of the embryo, and does not kick off until spring. Pregnancy then lasts about six weeks though it may again be slowed down if conditions turn bad. Long-eared bats are unusual among bats in that twins are not uncommon.

Whereas at least some nyctophilin species remain common (the lesser long-eared bat Nyctophilus geoffroyi is found over most of Australia), others are rare or little-known. A species described from Lord Howe Island, N. howensis, is believed to be extinct. The most remarkable case of obscurity is Pharotis imogene which was not recorded between 1890 and 2012, over 120 years. Evidence of extreme rarity? Quite probably, but also possibly evidence of just how few people are paying attention to bats.

REFERENCES

Hall, L. S., & D. P. Woodside. 1989. Vespertilionidae. In: D. W. Walton, & B. J. Richardson (eds) Fauna of Australia vol. 1B. Mammalia pp. 871–886. Australian Government Publishing Service: Canberra.

Miller, G. S., Jr. 1907. The families and genera of bats. Smithsonian Institution, United States National Museum, Bulletin 57: i–xvii, 1–282, pls 281–214.

The Font of the Placentals

The large-scale incorporation of molecular data into phylogenetics over the last few decades has caused a revolution in our understanding of life's evolution. Taxa whose interrelationships were previously regarded as intractable have been opened up to study, and many of our previous views on relationships have been forced to shift. Because conflict always makes for a good story, certain cases of the latter have become causes celebres, receiving extensive attention in both the technical and popular literature. One of these subjects of particular interest, not surprisingly, involves the relationships of the living orders of mammals.

Reconstruction of Arctostylops steini by Brian Regal, from Janis et al. (1998). The arctostylopids are a Palaeocene to Eocene group of mammals of uncertain affinities but probably belonging somewhere in the Boreoeutheria.

A lot of this attention has focused around the revelation of the Afrotheria, a grouping of animals (tenrecs, elephant shrews, hyraxes, aardvarks, elephants and manatees) with likely African origins that was completely unsuspected by studies based on morphological data only but which molecular studies have identified with ever-increasing levels of support. Recent molecular studies of placental phylogeny have agreed on three basal divisions within the placental mammals: the Afrotheria, the Xenarthra (armadillos, anteaters and sloths, a grouping that was recognised even before the advent of molecular data), and the remaining placentals in the largest of the three, the Boreoeutheria.

To the best of my knowledge, the Boreoeutheria is a clade that has also so far been supported by molecular data only with no morphological features yet recognised as defining the group. Nevertheless, its support can be considered as well established. The name Boreoeutheria refers to the clade's likely northern origins in contrast to the more southern distribution of the other two. Within the Boreoeutheria, molecular studies indicate a basal divide between the Euarchontaglires on one side and the Laurasiatheria on the other. The Euarchontaglires include the primates and rodents (as well as a handful of smaller orders). The Laurasiatheria include the Eulipotyphla, a group of insectivorous mammals including shrews, moles and hedgehogs, as sister to a clade containing bats, carnivorans, perissodactyls and artiodactyls.

Molecular phylogeny of mammals, from Springer et al. (2004) (note that not all branches shown in this tree are supported by all studies).

This all has interesting ramifications for the early evolution of placentals. There is an extensive fossil record of mammals from the Palaeocene, the epoch of time immediately following the end of the Cretaceous. However, most of these mammals do not belong to the orders alive today and their exact relationships to living mammals remain open to debate. The molecule-induced shake-up of pacental relationships just increased this uncertainty: for instance, the interpretation of a given group of fossil mammals as close to the common ancestry of perissodactyls and elephants rather goes out the window when perissodactyls and elephants are no longer thought to be closely related. And detailed studies that may resolve these issues remain few and far between. One of the most notable analyses in recent years has been that by Halliday et al. (2017) which covered most of the well-preserved placentals and their close relatives from the Cretaceous and Palaeocene periods. However, it is difficult to say just what to make of their results. The unconstrained analysis of their data presents results that remain deeply inconsistent with the molecular tree. Conversely, constraining the analysis to more closely match the molecular data provides results that are intriguing but difficult to accept at face value; I suspect they may be artefacts of the algorithm forcing taxa into the least unacceptable position for inadequate data. Suggesting that pangolins are the last specialised survivors of a broad clade of condylarths, pantodonts, notoungulates and creodonts is... I suppose not a priori impossible, but definitely a big call. A later analysis based on an expanded version of the same data set by Halliday et al. (2019) irons out some of the kinks but still fails to resolve the base of the Boreoeutheria beyond a massive polytomy of 25 branches (an icosipentatomy?). The Euarchontaglires are recovered as a clade but not the Laurasiatheria or any of its molecular subgroups above the ordinal level. And while some of the newer analysis' placements may seem like an improvement (notoungulates are placed as the sister to litopterns instead of hanging out with pangolins), others may still raise an eyebrow (mesonychids are associated with carnivorans but viverravids and miacids are not).

As always, the best answer to this conundrum is likely to involve more research. While researching this post, I did come across comments from people suggesting issues with the Halliday et al. data. Frankly, for a data set of this size (involving 248 taxa and 748 characters in the 2019 paper), it would be incredible were it otherwise. I know from my own experience that as you add more characters and taxa to a phylogenetic analyses, the challenge of keeping everything in line rises exponentially, and the data sets I've dealt with have been nowhere near the size of this one. Nevertheless, it's a start. And we can but hope that even those who find fault with it ultimately take it as inspiration to themselves do better.

REFERENCES

Halliday, T. J. D., M. dos Reis, A. U. Tamuri, H. Ferguson-Gow, Z. Yang & A. Goswami. 2019. Rapid morphological evolution in placental mammals post-dates the origin of the crown group. Proceedings of the Royal Society of London Series B—Biological Sciences 286: 20182418.

Halliday, T. J. D., P. Upchurch & A. Goswami. 2017. Resolving the relationships of Paleocene placental mammals. Biological Reviews 92 (1): 521–550.

The Grisons

Spend a bit of time following discussions of nature documentaries and other popular representations of biodiversity, and one topic you're likely to see come up is the biases that tend to exist in what gets represented. Images from eastern and southern Africa predominate while the west and north of that continent get overlooked. Europe and North America receive much more attention than the temperate regions of Asia. Another region whose diversity tends to go underrepresented is South America. The casual observer might think this continent is all monkeys and jaguars but South America is also home to notable radiations of dogs, deer, rodents, and other animals that many people would associate more with other parts of the world. Among these overlooked elements of the South American fauna are the local species of mustelid, including the grisons of the genus Galictis.

Greater grison Galictis vittata, copyright Tony Hisgett.

Grisons are somewhat ferret- or skunk-like animals found across almost the entirety of South America, and north into southern Mexico. They are greyish in colour dorsally (the name 'grison' itself means 'grey') with a black face and underparts. A pale stripe separates the upper and lower parts across the top of the face and continues diagonally back to the shoulders. They feed on small vertebrates and tend to be solitary hunters though they may sometimes form small family groups. They are primarily terrestrial and diurnal in habits. They have a reputation for ferocity; residents of Chile apparently have a history of using comparisons to grisons to describe unchecked rage (Yensen & Tarifa 2003b), in a similar manner to references to wolverines and honey badgers in other parts of the world. Contrasting colour patterns like those of the grisons are associated in other musteloids (such as skunks) with the production of offensive odours for defence, and grisons also produce strong-smelling secretions from their anal glands. Though some sources have claimed the odour produced by the lesser grison to be worse than a skunk's, it appears that these reports are exaggerated (Yensen & Tarifa 2003b).

Lesser grison Galictis cuja, copyright Ken Erickson.

Most authors have recognised two species of grison, the greater grison Galictis vittata and the lesser grison G. cuja*, as corroborated by a recent taxonomic study of the genus by Bornholdt et al. (2013). As their names indicate, the greater grison is generally larger and more robust than the lesser, being about 60 to 76 cm in total length versus 44 to 68 cm for the lesser grison (Yensen & Tarifa 2003b). The tail is also proportionately shorter in the greater grison (30% of the total length for the greater, 40% for the shorter). Fur is relatively longer and denser in the lesser grison, giving it more of a fluffy look. Whereas the dorsal fur is always a plain grey in the greater grison, it may often have a yellowish tinge in the lesser (not always, though). The two are generally distinct in range and habitat, as well. The greater grison is an animal of tropical forests and inhabits the northern part of the genus' range in Central America and northern and western South America. The lesser grison inhabits drier habitats, in arid or temperate regions, and so occupies the southern and eastern parts of the continent. The ranges of the species are known to overlap in Bolivian and Paraguay where their respective biomes approach each other.

*Some sources have listed a third species G. allamandi but this seems have been something of a 'ghost' taxon born from confusion whether the name 'G. vittata' applied to the greater or lesser species.

The genus Galictis arrived in South America as part of the Great American Biotic Interchange, about three million years ago. The general consensus is that it is derived from the genus Trigonictis of the North American Pliocene. Indeed, it has even been suggested that the two North American species of Trigonictis might represent independent ancestors of Galictis, with the larger T. macrodon giving rise to the greater grison and the smaller T. cookii birthing the lesser grison (Yensen & Tarifa 2003a). This certainly would seem overly complicated, though, and molecular data are more in line with a more recent separation of the species.

REFERENCES

Bornholdt, R., K. Helgen, K.-P. Koepfli, L. Oliveira, M. Lucherini & E. Eizirik. 2013. Taxonomic revision of the genus Galictis (Carnivora: Mustelidae): species delimitation, morphological diagnosis, and refined mapping of geographical distribution. Zoological Journal of the Linnean Society 167: 449–472.

Yensen, E., & T. Tarifa. 2003a. Galictis vittata. Mammalian Species 727: 1–8.

Yensen, E., & T. Tarifa. 2003b. Galictis cuja. Mammalian Species 728: 1–8.

Predators of the European Eocene

Among mammals in today's modern fauna, the role of terrestrial carnivore is dominated by members of one particular lineage, known (appropriately enough) as the Carnivora. But travel back in time to the Eocene period, roughly 56 to 34 million years ago, and you'll find a range of now extinct groups sharing that role. This post is looking at one of those groups, the proviverrines.

The Proviverrinae are a subgroup of the Hyaenodontidae, one of the two families of carnivores commonly associated as the creodonts. I've discussed creodonts before, and the overhanging question of whether they form a coherent evolutionary group. Currently, my impression is that most mammal palaeontologists seem inclined to think that hyaenodontids and oxyaenids probably do not share an immediate common ancestry. However, nor is there any clear idea of what else either group may relate to.

Skull of Cynohyaenodon cayluxi, photographed by Ghedoghedo.

Historically, proviverrines have been treated as the basal grade from which other groups of hyaenodontids were derived with representatives known from Europe and North America. However, a phylogenetic analysis of early hyaenodontids by Solé (2013) lead to a division of the 'proviverrines' between three monophyletic subfamilies: the Proviverrinae proper, the Sinopinae and the Arfiinae. Under this system, the Proviverrinae are a uniquely European group. As is standard in mammalian palaeontology, proviverrines (in the strict sense) are distinguished from other hyaenodontids by features of the teeth. Notable among these is the presence of a double root on the first lower premolar of most proviverrines; other hyaenodontids have a single root on this tooth.

The earliest proviverrines are known from the very beginning of the Eocene (Solé et al. 2014). Current thinking is that their ancestors probably immigrated into Europe around this time from Africa. The Late Paleocene Tinerhodon disputatum from northern Africa resembles a proviverrine in overall appearance but was probably more basally placed in respect to hyaenodontids as a whole. The name 'Proviverra' can be read as 'early civet' and while proviverrines were not related to modern civets (which are, of course, true carnivorans) this is probably not a bad indication of the overall appearance of their original appearance. These were very small animals, probably less than 100 g in body weight, and probably had a fairly generalised diet of small vertebrates and invertebrates. At first, proviverrines seem to have been restricted to southern Europe, what is now Spain and the very southernmost part of France. Northern Europe was inhabited by the Arfiinae and Sinopinae, as well as species of Oxyaenidae (the other 'creodont' family). Sinopinae were also found in southern Europe and may have excluded the proviverrines from evolving larger size. However, the other hyaenodontids and oxyaenids went extinct in Europe not to long after the beginning of the Eocene. A turnover in the mammalian fauna of North America around this time appears to be due to a cooling of the climate; though the evidence for climate cooling is less clear in Europe, it seems reasonable that it was going through similar changes. With their competitors out of the picture, the proviverrines rapidly diversified into the regions and niches that had been left unoccupied.

Lesmesodon edingeri, photographed by Ghedoghedo.

The largest proviverrines, members of the genera Prodissopsalis, Paenoxyaenoides and Matthodon, would eventually reach weights of close to twenty kilograms, about as large as a medium-sized dog. They would also diversify in their habits. Members of the genera Oxyaenoides and Paenoxyaenoides were cursorial hypercarnivores, their dentition specialised for a diet almost exclusively of meat*, like that of a modern cat. Matthodon and Quercytherium, in contrast, were genera whose dentition showed more adaptations for cracking hard materials such as bone. They may have had lifestyles more like those of hyaenas, with Matthodon (which combined adaptations for hypercarnivory and bone-cracking) perhaps being more of an active hunter than Quercytherium.

*These two genera also provide an excellent example of the role of convergent evolution in the evolution of mammalian carnivores. Their appearance to other hypercarnivorous hyaenodontids was such that it was only recently that they were recognised as proviverrines rather than members of other subfamilies no longer thought to have been found in Europe. And not only are they remarkably convergent on other subfamilies, the phylogenetic analysis of proviverrines by Solé et al. (2014) suggests that they're not even directly related to each other within that clade.

Proviverrines remained the dominant mammalian carnivores in Europe for about the next twenty million years but then went into a sharp decline. This reversal of fortunes may have been due to the increasingly cool, dry conditions developing at this time, and/or it may have been related to competition from the first true carnivorans arriving in Europe. The larger, more specialised proviverrines disappeared rapidly when their time came. The last surviving genus, Allopterodon, was a small form, little more than one kilogram in weight, and had a generalised dentition indicating a relatively unspecialised diet. This may have been a return to something like the lineage's original form but it would not save it: by the end of the Eocene, the proviverrines would be completely extinct.

REFERENCES

Solé, F. 2013. New proviverrine genus from the Early Eocene of Europe and the first phylogeny of Late Palaeocene–Middle Eocene hyaenodontidans (Mammalia). Journal of Systematic Palaeontology 11 (4): 375–398.

Solé, F., J. Falconnet & L. Yves. 2014. New proviverrines (Hyaenodontida) from the early Eocene of Europe; phylogeny and ecological evolution of the Proviverrinae. Zoological Journal of the Linnean Society 171: 878–917.

The Cervini: Deer of Temperate Eurasia (and Beyond)

A couple of years back, I presented you with a post giving a quick overview of the classification of deer. For this post, I'm going to look a bit closer at a particular subgroup of deer: the species of the tribe Cervini.

Wapiti Cervus canadensis, photographed by Mongo.

For most people outside the Americas, a member of the Cervini will probably represent the first image that comes to mind when picturing a deer. The same goes for many Americans, for that matter, though in that part of the world they face a bit more competition. Cervins are the most diverse group of deer in temperate Eurasia, with representatives also being found in northernmost Africa, North America, India and southeast Asia (as well as introduced species in Australasia). The Monarch of the Glen was a cervin: specifically, a red deer Cervus elaphus. Bambi, in his Disney film incarnation, was also a cervin, a wapiti C. canadensis (in his original literary form, probably less familiar to modern audiences with little interest in Austrian novels about all the miserable ways that animals can die, he was a roe deer Capreolus capreolus and so not a cervin) (Edit: Scratch that, he's a apparently a non-cervin white-tailed deer, see comment below). The group has long been recognised by features of the skull and leg bones, and also is well supported by molecular data (Heckeberg 2020). Males produce antlers with multiple branches (at least in typical individuals) with the branches or tines usually directed forwards from the main shaft of the antler (the Père David's deer Elaphurus davidianus differs from other cervins in having the tines directed rearwards). The first of these branches, the brow tine, usually originates close to the base of the antler. In a number of Asian species, such as the chital Axis axis and sambar Cervus unicolor, there is usually on one more branch on the antler so each antler ends with three points. Species with such antlers are generally found in dense forests and their simpler antlers may represent an adaptation to these habitats (Heckeberg 2020). Other cervin species may have more extensively branched antlers with a tendency for antler complexity to correlate with overall body size; the largest living cervins, the red deer and wapiti, also have the most branched antlers. Larger extinct species had even more extravagant headgear with the apex of insanity being perhaps the bush-antlered deer Eucladoceros dicranios of the lower Pleistocene of Europe: each antler of this species might carry a dozen points.

Skull of Eucladoceros dicranios, photographed by Aldo Cavini Benedetti.

To describe the classification of cervins as having recently been in a state of flux is something of an understatement. A conservative presentation of the group may refer to thirteen or fourteen living species in four genera (e.g. Macdonald 1984). More recent authors, however, might refer to up to ten genera and nearly forty species. In a way, this difference is not really as dramatic as it may seem: multiple subspecies have long been recognised for most cervin species and some authors have argued for the recognition of many of these 'subspecies' as distinct species. Classification at generic level has mostly been affected by recognition that the genus Cervus as previously recognised is not monophyletic. Most recent authors agree on the recognition of at least four genera of Cervini (Cervus, Dama, Axis and Rucervus) with two further genera (Rusa and Elaphurus) also commonly recognised.

Persian fallow deer Dama mesopotamica, copyright Rufus46.

The genus Dama is usually recognised as including two species, the fallow deer D. dama and Persian fallow deer D. mesopotamica. These species are readily distinguished from other cervins by the form of their antlers which are distally palmate. Palmate antlers are also characteristic of the extinct giant Irish elk Megaloceros giganteus and many recent authors have regarded the two as closely related. The white spots that many deer species possess when young are commonly retained by fallow deer into adulthood though the coat will often become darker and the spots disappear during winter. Melanistic and leucistic individuals of fallow deer are also common. Defining the native range of the fallow deer is a bit of a tricky question. This inhabitant of open woodlands is currently widespread in Europe but was probably restricted to a region of the eastern Mediterranean during the last ice age. Its current range in northern Europe may in large part be the result of human transportation. The fallow deer has also been widely introduced elsewhere: herds may now be found in numerous locations in Africa, Australasia, North and South America. The Persian fallow deer, in contrast, is now endangered, its range restricted to a small number of localities in Iran. Indeed, it was once thought to be extinct prior to the rediscovery of a population of about two dozen individuals in the mid-1950s; the current population is perhaps only a few hundred.

Thorold's deer Cervus albirostris, copyright Heather Paul.

The genus Cervus in its current, more restricted sense includes the red deer and wapiti as well as the sika C. nippon* of eastern Asia. Sika are generally smaller than the other two species and, like fallow deer, usually retain the juvenile spots into adulthood. Excluding occasional small accessory branches, the antlers of sika also possess no more than four tines (Heckeberg 2020) in contrast to the commonly further branched antlers of red deer and wapiti. Four-tined antlers are also characteristic of the Thorold's or white-lipped deer C. albirostris, an inhabitant of the Tibetan Plateau that has sometimes been treated recently as the only representative of a separate genus Przewalskium. White-lipped deers have broad, cow-like hooves for navigating the steep, rocky slopes of their homeland. More commonly accepted classification-wise is the separation of two species found in southern Asia, the rusa C. rusa and sambar C. unicolor, as the genus Rusa. Both these species have three-tined antlers and their fawns lack spots.

*Commonly referred to as the sika deer. 'Sika' (or, as it's more commonly transliterated these days, 'shika') is Japanese for deer, so the common vernacular name of Cervus nippon is, indeed, 'deer deer'. The same issue arises for the rusa deer in Malay.

Chitals Axis axis, copyright Charles J. Sharp.

Axis is a genus of four species of smaller forest-dwelling deer found in southern Asia. Antlers are generally three-tined with the upper beams curving inwards towards each other. The chital remains spotted at maturity whereas the other species loose their spots. These species include the hog deer A. porcinus, named for its low, short-legged build, and two closely related insular species. Recent years have seen some authors separate the hog deers as a separate genus Hyelaphus, restricting Axis to the chital, owing to molecular phylogenies casting doubt on the genus' monophyly. However, it seems that these studies may have been mislead by a contaminated sample for the hog deer (Gilbert et al. 2006) and other studies have retained a monophyletic Axis. The thamin Rucervus eldii and barasingha R. duvauceli are also found in southern Asia where they tend to be associated with marshy habitats. Their antlers curve outwards then inwards to form a bow-shaped curve; those of the thamin are three-tined whereas the barasingha possesses further tines, sometimes up to ten on each antler. Again, some studies have questioned the monophyly of Rucervus and suggested the thamin be moved to a separate genus Panolia.

Père David's deers Elaphurus davidianus, copyright Peter O'Connor.

Finally, there is Père David's deer, arguably the weirdest of all the cervins, most often placed in its own genus Elaphurus but sometimes included in Cervus. By the time this species became known to European naturalists, it was already extinct in the wild, surviving only as a herd kept in a hunting garden near Peking belonging to the emperor of China. This herd was exterminated during the Boxer Rebellion but specimens that had been transported to Europe saved the species from total extinction. It is now widely kept in captive herds and has also been returned to the wild in a couple of locations in China. Père David's deer has a number of features that make it stand out from other deer: as well as the aforementioned backwards antlers, it has wide, splayed hooves and a remarkably long tail. But in other regards, Père David's deer is not anywhere as weird as it should be. In particular, its karyotype is very similar to that of the red deer: close enough, in fact, that not only are the two species capable of hybridising in captivity but the resulting hybrids are fully fertile (such matings are unlikely in the wild owing to the two species normally having different breeding seasons). Heckeberg (2020) found that Père David's deer was associated with Cervus species in analyses of nuclear genes and cranial characters but with Rucervus species in analyses of mitochondrial genes and dentition; other authors had previously found similar results. It has been suggested that these schizoid tendencies with regard to phylogenetic analysis might indicate a hybrid origin for Père David's deer from ancestors related to the wapiti on one side and the thamin on the other. Such a hybridisation event would have happened some time ago—fossils related to Père David's deer seem to date back at least to the late Pliocene—allowing enough time to pass for the new population to develop its own idiosyncracies not acquired directly from either parent.

Variations on a Tayra

Subspecies can be a funny thing in the world of animal taxonomy. Millions of litres of ink have been spilt over the years arguing over how one defines a species but a lot less has been invested in discussing the nature of subspecies. For some popular species concepts (such as the most popular iteration of the 'phylogenetic species concept'), one might question whether any concept of subspecies could be applied at all (I could suggest some hypothetical situations but just how applicable or practical they are is a further matter). Essentially, most subspecies concepts distill down to 'a population that is distinct enough to warrant recognition but somehow doesn't quite qualify as a species'. Historically, the rank has tended not to receive a lot of usage among animals outside groups subject to particularly high levels of taxonomic attention—most particularly, vertebrates and butterflies—and many currently recognised animal subspecies were first named in days when taxon descriptions tended to be much briefer and taxonomists were under less pressure to explain their reasoning. Because subspecies tend to be, by their nature, vague and difficult to define, and because evaluating them often requires detailed population analysis within a species, these historical subspecies have a tendency to linger, unchallenged, in taxonomic listings. And with that as background, tayras.

Tayra Eira barbara photographed in Peru, copyright eMammal. Photography location would indicate this individual to be either E. b. madeirensis or E. b. peruana.

The tayra Eira barbara is a large mustelid (a member of the family including weasels, otters and badgers) found in warmer regions of Central and South America, its distribution extending down to about the level of the southern edge of Brazil. They are long-bodied but robust animals, kind of looking like a 'roided-up stoat. They grow to a head-body length of two feet or more (up to about 71 centimetres) with a tail about two-thirds as long again. Adult males tend to be a third as large again as females and more muscular around the fore quarters. Comparisons have often been made between tayras and the martens Martes of the Northern Hemisphere and molecular studies confirm a relationship between these two genera, as well as the wolverines Gulo. Closer fossil relatives are known from North America and it seems likely that the tayra originated on that continent then spread southwards. Ruiz-García et al. (2013) suggested that the degree of genetic divergence between tayras found in South America might indicate the species may have arrived there about eight million years ago, before the formation of the Panamanian land bridge. Tayras are not the only species for which this possibility has been suggested; these early arrivals may have reached South America by island-hopping between earlier-emerging segments of the eventual connection.

Tayras are diurnal omnivores, their known diet ranging from fruits to small animals to honey. In captivity, it seems they will accept pretty much anything offered to them. Tayras are the only animals other than humans that have been recorded caching unripe fruit in order to eat it after it finishes ripening. It is still not certain to what degree tayras are solitary or social; though commonly regarded as solitary, they have been recorded hunting howler monkeys in groups (Shostell & Ruiz-Garcia 2013). Tayras are mostly found in forests; in some areas they may adjust to more open habitats but seemingly only under sufferance (Presley 2000). Though not regarded as 'arboreal' per se, tayras are adept climbers. Their well developed carpal vibrissae ('whiskers' on the wrists) presumably contribute to this ability. Their wide distribution and adaptability mean that tayras are not currently regarded as of conservation concern though habitat degradation has reduced their numbers in some areas.

Tayra from Belize, presumably the light-headed Eira barbara senex, from Wikimedia Commons.

The body and tail of tayras are generally dark brown or black with the head being distinctly lighter in coloration (light brown or grey to yellow). Leucistic and albino individuals are not that uncommon (yellow tayras are apparently particularly common in Guyana). A patch of pale coloration, varying from a spot to a broad triangle, is often (but not always) present on the chest and throat. Recent taxonomic listings (e.g. Presley 2000) have recognised seven subspecies of tayra distinguished by coloration. The Mexican Eira barbara senex has a greyish white head with the light coloration extending to dark yellow shoulders and a dark brown body. Eira barbara inserta, found in southern Honduras and Nicaragua, is a dark subspecies with a dark brown head, black body and no throat patch. The Colombian E. b. sinuensis is darker than E. b. senex with the nape a darker brown than the head; it may or may not possess a throat patch. Eira barbara barbara, found in southern Brazil, eastern Bolivia and Paraguay, is lighter than E. b. sinuensis but darker than E. b. senex and has a yellowish throat patch. The northern Brazilian E. b. madeirensis is a chocolate brown with the head slightly lighter than the body; again, a throat patch may or may not be present. The Peruvian and western Bolivian E. b. peruana is similar to the last subspecies but has darker legs and a black tail. Finally, E. b. poliocephala, which has a distribution centred on the Guianas, is similar to E. b. barbara but with a darker yellow throat patch and yellow shoulder patches that sometimes merge with the throat patch to form a complete collar.

Tayra photographed in a zoo in Panama, copyright Dirk van der Made. Being a zoo individual, its origins are a bit more open than the other individuals shown on this page, but Panama is home to Eira barbara inserta and E. b. sinuensis.

Such is the received wisdom as recorded by Presley (2000) but does it accurately reflect population distributions? Ruiz-García et al. (2013) conducted an analysis of mitochondrial genes from tayras representing the five South American subspecies (i.e. excluding E. b. senex and E. b. inserta). They found that of these five subspecies, only E. b. poliocephala (as represented by specimens from French Guiana) could potentially be differentiated genetically. Samples from the ranges of the other four 'subspecies' were intermingled in analyses, leading Ruiz-García et al. to suggest that they should be merged into a single subspecies E. b. barbara (it may also be worth me mentioning that, when I was looking for images to illustrate this post, I had difficulty finding ones in which the supposed differences between subspecies were recognisable). Of course, that leaves the status of the two Central American subspecies undetermined. It may be of note that they seem to be more distinct in appearance than some of the hitherto-recognised South American subspecies but it remains to be seen just how significant this is.

REFERENCES

Presley, S. J. 2000. Eira barbara. Mammalian Species 636: 1–6.

Ruiz-García, M., N. Lichilín-Ortiz & M. F. Jaramillo. 2013. Molecular phylogenetics of two Neotropical carnivores, Potos flavus (Procyonidae) and Eira barbata (Mustelidae): no clear existence of putative morphological subspecies. In: Ruiz-Garcia, M., & J. M. Shostell (eds) Molecular Population Genetics, Evolutionary Biology and Biological Conservation of Neotropical Carnivores pp. 37–84. Nova Publishers: New York.

Shostell, J. M., & M. Ruiz-Garcia. 2013. An introduction to Neotropical carnivores. In: Ruiz-Garcia, M., & J. M. Shostell (eds) Molecular Population Genetics, Evolutionary Biology and Biological Conservation of Neotropical Carnivores pp. 1–34. Nova Publishers: New York.

The Mouse Shrews of Africa

Shrews are one of the less appreciated groups of mammals. Small (some are among the smallest mammals on earth), skulking, they are often overlooked but are nevertheless represented by a diversity of species in many parts of the world. Among this diversity are the mouse shrews of the genus Myosorex.

Forest shrew Myosorex varius, from Roberts (1951).

Myosorex is a genus of nearly twenty known species of shrew, some of which have only been identified very recently, with more probably yet to be described. They are known in the modern fauna only from sub-Saharan Africa, though the fossil record indicates they once extended as far north as Spain (Furió et al. 2007). Mouse shrews differ from most other living shrew genera (except the closely related Congosorex) in the presence of a tiny vestigial tooth in the lower jaw behind the first antemolar (the tooth behind the incisors, so called in shrews because it is unclear whether it corresponds to a canine or premolar in relation to the teeth of other mammals). Because their teeth lack the red pigment found in shrews of the subfamily Soricinae, Myosorex have historically been classified with the white-tooth shrews of the Crocidurinae. However, the presence of white teeth is, of course, a primitive feature of questionable significance phylogenetically. Instead, more recent authors have pointed to the retention of the second antemolar and other features to support recognising Myosorex and related African shrew genera as relictual members of a third subfamily, the Myosoricinae, that may also include a number of earlier fossil shrews (this group has also been known as the 'Crocidosoricinae' but 'Myosoricinae' is the name with priority; the argument by Furió et al., 2007, that the latter name cannot be used for the broader subfamily because it was originally used only for the African genera has no standing under current nomenclatorial rules).

Foraging forest shrew, copyright Johnny Wilson.

In the modern fauna, Myosorex species have a very scattered distribution. Species found in central and eastern Africa are restricted to high mountains, among moist, densely vegetated environments. Species found in southern Africa are often found in similar habitats. However, the species M. varius is also found in drier locations at lower altitudes, closer to the South African coast. Nevertheless, it is still restricted to areas with high seasonal rainfall, or (in part of Western Cape Province) zones dominated by succulent vegetation where low levels of actual rainfall may be compensated for by precipitation from mist (Meester 1958). The distribution of the genus as a whole is marked by a broad gap of over 1600 km separating the northern limit of species in South Africa and Zimbabwe from their nearest neighbours to the north in the DRC and Kenya, a gap that they were presumably only able to cross in the past when climate conditions were more amenable.

This sensitivity to environment means that Myosorex species may be very vulnerable to changes in habitat. Several species are restricted to limited ranges and several are recognised as potentially endangered. The prospect of climate change makes this vulnerability even worse: as levels of rainfall decrease, mouse shrews will be forced to retreat to ever higher altitudes, and there's only so high they can go before running out of mountain.

REFERENCES

Furió, M., A. Santos-Cubedo, R. Minwer-Barakat & J. Agustí. 2007. Evolutionary history of the African soricid Myosorex (Insectivora, Mammalia) out of Africa. Journal of Vertebrate Paleontology 27 (4): 1018–1032.

Meester, J. 1958. Variation in the shrew genus Myosorex in southern Africa. Journal of Mammalogy 39 (3): 325–339.

Fishing Mice

In a 1950 discussion of the origins of the fauna of South America, the great American palaeontologist G. G. Simpson dismissed the enormous radiation of muroid rodents in that continent as mere "field mice" exhibiting little regional differentiation. George Gaylord Simpson may have been one of the leading thinkers in mid-20th Century evolutionary theory, but in this respect he was just plain wrong. The South American mice and rats include a wide variety of divergent forms, some of them specialised in surprising ways. Consider, for instance, the fishing mice of the Ichthyomyini.

Illustration of Stolzmann's crab-eating mouse Ichthyomys stolzmanni by Joseph Smit.

The Ichthyomyini are a small assemblage of less than twenty species of mice found between Mexico and the north of South America from Peru to French Guiana (Voss 1988). Though some species are known from lower altitudes, the majority are found in alpine habitats in association with fast-flowing mountain streams (albeit in no location are they known to be common). Ichthyomyins seem to show a particular preference for hanging around waterfalls (Barnett 1997) and are not found in association with standing water such as swamps or ponds. They are moderate in size, ranging from ten to twenty centimetres in length excluding the tail. They show a number of adaptations for foraging underwater: the hind feet are partially webbed and have a more or less elongate fringe of stiff hairs that aid in swimming, the tail is furry rather than scaly, the eyes are small, the external ears are reduced in size (in a couple of species they are completely hidden by the fur and in one, Anotomys leander, the external pinnae are missing entirely), and the whiskers are long, strong, and arranged in such a way that they almost look more like the whiskers of a sea lion than of a mouse. The nerves associated with these whiskers are also enlarged and they evidently provide the main means of finding food.

Peruvian fish-eating rat Neusticomys peruviensis, copyright Carlos Boada.

Or, as I should say, finding prey. As far as we know, these mice seem to be entirely carnivorous. Only a couple of examples are known of specimens with plant matter in their stomachs and the significance of those finds remains uncertain. The primary source of food for most species is small invertebrates such as aquatic insects. Where freshwater crabs are available, a number of species show preferences for those. In larger species, the diet may be supplemented to a greater or lesser degree by small vertebrates such as fish or tadpoles. In line with their carnivorous diet, ichthyomyins are also characterised by a shorter, less complicated gut than other mice. Little is known about breeding and nesting habits in ichthyomyins. A specimen of Chibchanomys kept in captivity made tunnels in mossy vegetation (Barnett 1997). The few known specimens of gravid females indicate that litters are small with no more than two foetuses being carried at a time.

Voss (1988) recognised five genera of Ichthyomyini. The largest of these, Neusticomys, includes about half a dozen species that may more closely resemble the ancestral morphology for the group. Their hind feet are narrower than those of other ichthyomyins and the fringe of swimming hairs is shorter (Packer & Lee 2007). Where one species found in Colombia and Ecuador, Neusticomys monticolus, overlaps in range with Anotomys leander, it shows a preference for more sheltered sections of stream banks whereas A. leander is found in more exposed rapids.

Undescribed species of Chibchanomys, copyright Alexander Pari.

In most ichthyomyins, the coat consists of a layer of dense, woolly underfur covered by an overcoat of long guard hairs mixed with glossy, often distally flattened awn hairs. In Anotomys leander, Chibchanomys trichotis, and Neusticomys monticolus, the awn hairs are missing so these species have a dull grayish black appearance overall rather than than the glossy coat of other species. Chibchanomys trichotis retains minute external ear flaps albeit not ones that are visible past the coat; Anotomys leander, as noted above, lacks external ear flaps but does have the positions of the ear openings marked by a prominent white spot. Both these last two species were placed in monotypic genera by Voss (1988), but Barnett (1997) refers to an at-that-point undescribed species of Chibchanomys.

The remaining two genera were recognised by Voss (1988) as including four species apiece. Species of Rheomys, found in the mountains of Central America, have the most extensively webbed hind feet among the ichthyomyins. This is the only genus of fishing mice found in Central America; the other genera are all restricted to South America. The genus Ichthyomys includes the largest species of the group and also the species that feed on the highest proportion of vertebrates. This difference in diet is reflected in their dentition: Ichthyomys species have proportionately larger incisors and smaller molars than other ichthyomyins, with greater emphasis on using the incisors to grasp and slice struggling prey.

Rheomys raptor, from Villalobos-Chaves et al. (2016).

All told, the ichthyomyins are a remarkable radiation. Ecologically, they are close parallels to forms found elsewhere such as water shrews or desmans, but most other semi-aquatic mammals are distinctly larger in size. Even with less than twenty species, the ichthyomyins represent more species than there are of similarly sized semi-aquatic mammals anywhere else in the world. However, as noted above, ichthyomyins are not common anywhere they occur, and factors such as deforestation and climate change could represent a significant threat to their survival. It would be unfortunate if this remarkable radiation was to fade away.

REFERENCES

Barnett, A. A. 1997. The ecology and natural history of a fishing mouse Chibchanomys spec. nov. (Ichthyomyini: Muridae) from the Andes of southern Ecuador. Zeitschrift für Säugetierkunde 62: 43–52.

Packer, J. B., & T. E. Lee Jr. 2007. Neusticomys monticolus. Mammalian Species 805: 1–3.

Voss, R. S. 1988. Systematics and ecology of ichthyomyine rodents (Muroidea): patterns of morphological evolution in a small adaptive radiation. Bulletin of the American Museum of Natural History 188 (2): 259–493.

Guenons

The common perception of monkeys tends to be dominated by a relatively small number of species, generally those most commonly seen in zoos, such as capuchins, macaques, baboons or tamarins. But as is usual when it comes to biodiversity, there are a lot of varieties of monkey out there that may be less familiar to the general public. This post will look at one of those less familiar groups: the guenons of the genus Cercopithecus.

Moustached monkey Cercopithecus cephus, copyright Rufus46.

Cercopithecus is a genus of monkeys found in sub-Saharan Africa. The exact number of species has shifted around a bit (though it currently sits around twenty). Some authors have included almost all species of the monkey tribe Cercopithecini, characterised by self-sharpening lower incisors and four cusps on the lower third molars (Lo Bianco et al. 2017), in the single genus Cercopithecus. However, more recent authors have tended to favour dividing this tribe between a number of phylogenetically and ecologically distinct genera. Under this latter system, Cercopithecus would be restricted to a group of more arboreal species. A number of these species have been divided between multiple subspecies and there may be some back and forthing about what is recognised as which. One entirely new species, previously not even known as a subspecies, was described as recently as 2012 by Hart et al.: the lesula C. lomamiensis.

Young female lesula Cercopithecus lomamiensis, from Hart et al. (2012).

A large part of this uncertainty relates to the fact that Cercopithecus species are most diverse in dense forests of western and central Africa, in regions that may be both physically and politically difficult to access and which have received less attention from researchers than others. The aforementioned lesula was described from the Lomami River basin near the middle of the Democratic Republic of the Congo (the one that used to be called Zaire, though I think they prefer not to talk about it). Another Congolese species, the dryas monkey C. dryas, was long thought to be known from only a single juvenile specimen until it was realised that the adult form had been described as a separate species C. salongo. It's still only known from a handful of records and is thought to be critically endangered.

Diana monkey Cercopithecus diana, copyright Ikmo-ned.

Some species of guenon are notable for their striking colour patterns. Perhaps the species I've most commonly seen in zoos is the diana monkey C. diana, native to the region between Sierra Leone* and the Côte d'Ivoire (though it is possible that at least some of these 'diana monkeys' were actually roloway monkeys C. roloway, until recently treated as a subspecies of the diana monkey). This species has a bright white throat, chest and front of the fore arms that contrasts with the black face and dark grey back. It also has a white band across its brow which is where its name comes from, the band having been thought to resemble the crescent moon. De Brazza's monkey C. neglectus of central Africa has a crescent-shaped orange mark on its forehead and a white muzzle and beard, making it look reminiscent of a grumpy old man (Wikipedia claims that it has also been dubbed the 'Ayatollah monkey'). Male De Brazza's monkeys also have a bright blue scrotum. Large bright blue patches are also present around the scrotum and backside of males in the lesula and the owl-faced monkey C. hamlyni.

*Having grown up in New Zealand in the 1980s, I'm going to have that stuck in my head all day now. Nothing to do with the subject of this post, I just thought I'd mention it.

Male De Brazza's monkey Cercopithecus neglectus, copyright Heather Paul.

Guenons tend to be found living in small troops consisting of one adult male and a harem of females with their offspring; unmated adult males will be found living solitary lives. Males are usually larger than females, up to about 1.5 times the size of their mates. Multiple guenon species may be found in a single location though closely related species tend not to overlap. Famously, hybrids have been described from the Kibale forest in Uganda between the blue monkey C. mitis and the red-tailed monkey C. ascanius, two species that are quite distinct in external appearance. Larger species such as the spot-nosed monkey C. nictitans and the blue monkey tend to eat a higher proportion of leaves in their diet. Smaller species such as the mona monkey C. mona may be more insectivorous (Macdonald 1984).

Blue monkeys Cercopithecus mitis stuhlmanni, copyright Charles J. Sharp.

The origins of the Cercopithecus radiation are relatively recent with the tribe Cercopithecini as a whole probably originating in the late Miocene (Lo Bianco et al. 2017). Karyological studies of the group show a wide variation in chromosome number from 58 in the diana monkey to 72 in the blue monkey. In contrast, the sister group of the Cercopithecini, the Papionini (which includes baboons and macaques) always has 42 chromosomes. Polymorphism in chromosome arrangements has also been described within Cercopithecus species. The possibility that this gene variability is related to their rate of speciation remains a worthwhile line of study.

REFERENCES

Hart, J. A., K. M. Detwiler, C. C. Gilbert, A. S. Burrell, J. L. Fuller, M. Emetshu, T. B. Hart, A. Vosper, E. J. Sargis & A. J. Tosi. 2012. Lesula: a new species of Cercopithecus monkey endemic to the Democratic Republic of Congo and implications for conservation of Congo's central basin. PLoS One 7 (9): e44271.

Lo Bianco, S., J. C. Masters & L. Sineo. 2017. The evolution of the Cercopithecini: a (post)modern synthesis. Evolutionary Anthropology 26: 336–349.

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

Moustached Bats, Ghost-faced Bats

In the theatre of mammal diversity, there are two groups that loom above all their competitors. The most diverse of the generally recognised mammalian orders, by a healthy margin, is the rodents. Nevertheless, they are still given a good run for their money by the silver medalist, the bats. There is somewhere in the region of 1100 known living species of bat, a number that has continued to increase in recent years as study progresses. This post will focus on one particular group of bats, the Mormoopidae.

Ghost-faced bat Mormoops megalophylla, copyright Merlin Tuttle.

Mormoopids are a group of bats found in warmer parts of the Americas. They commonly go by the names of moustached bats or funnel-eared bats, at least to the extent that any type of bat can be said to 'commonly' go by anything. They are fast-flying, insectivorous bats that roost in colonies in hot, humid caves. These colonies can be sizable: at least one colony of Wagner's moustached bat Pteronotus personatus was estimated to include more than 16,000 individuals (de la Torre & Medellín 2010). In the United States, mormoopids are currently restricted to the south-west (in the form of the ghost-faced bat Mormoops megalophylla) but subfossil from Florida indicate a wider distribution in the past (Simmons & Conway 2001).

Mormoopids belong to the Noctilionoidea, a distinctly Neotropical group of bats that also includes the leaf-nosed bats of the Phyllostomidae and the Noctilio bulldog bats (and also possibly the short-tailed bat Mystacina of New Zealand, because why make things simple?) The biggest difference between mormoopids and other noctilionoids lies in the structure of their shoulders. In most bats, the trochiter (one of the tubercles at the top of the humerus) is enlarged to form a secondary articulation with the scapula. This strengthens the shoulder joint, presumably allowing the production of more power for flight. Mormoopids, however, lack this enlarged trochiter. I must confess to being unsure just what is the significance of this alteration; mormoopids remain fast fliers (de la Torre & Medellín 2010). Are they perhaps sacrificing a bit of endurance for the sake of higher mobility?

Wagner's moustached bat Pteronotus personatus, copyright Bernard Dupont.

Current classifications of the mormoopids recognise two genera in the family, Mormoops and Pteronotus. Mormoops species have a shorter head than Pteronotus species (so short, in fact, that the braincase is wider than it is long), with a markedly upturned snout. Basically, they have a skull like a pug dog. In a revision of the family, Simmons & Conway (2001) recognised two living species of Mormoops and six of Pteronotus, plus an additional species of each described from subfossil remains from Cuba. Pteronotus was also divided between three subgenera. The type subgenus included two species, Pt. davyi and Pt. gymnonotus, known as naked-backed bats because the membrane for their wings attaches close to the spine so the body fur is not visible in dorsal view (in other mormoopids, the wings attach along the sides of the body). Three of the remaining species (Pt. personatus, the sooty moustached bat Pt. quadridens and Macleay's moustached bat Pt. macleayi) were placed in a morphologically generalised subgenus Chilonycteris. The remaining living species was Parnell's moustached bat Pt. parnellii, placed in its own subgenus Phyllodia.

Pteronotus parnellii was the only known mormoopid, and in fact the only Neotropical bat of any kind, to use high duty cycle echolocation. Echolocation, of course, works through the bat emitting calls and listening for when they bounce back from surrounding objects. The problem is that the noise produced while emitting calls can drown out returning echoes. As a result, most echolocating bats use what is called low duty cycle echolocation. Individual echolocation calls are spaced apart so the bat has time between each call to listen for echoes. High duty cycle echolocation is used by two Old World bat families, the Rhinolophidae and Hipposideridae, as well as Pteronotus parnellii. These bats have learnt the trick of emitting calls continuously and recognising returning echoes by their different frequency. This allows each bat to build up a more detailed picture of its surrounds, allowing for greater mobility in complex environments such as around dense forest.

Parnell's moustached bat Pteronotus parnellii, copyright Alex Borisenko.

In recent years, however, mormoopid systematics have been given a shake-up. Many of the mormoopid species recognised by Simmons & Conway (2001) could be divided between multiple subspecies. Recently, a molecular analysis of Pteronotus species by Pavan & Marroig (2016) found strong genetic divergence between most of these subspecies. As a result, they proposed raising the distinct subspecies to species level, effectively raising the number of living Pteronotus species from six to fifteen. Some of these species could also be separated on the basis of morphometric and acoustic data; others exhibited morphometric overlap but were geographically distinct. 'Pteronotus parnellii' was the most diverse, being divided into eight named species plus an unnamed population that may warrant species recognition. The question that this immediately raises: is the use of high duty cycle echolocation a feature of all nine of these species, or might it turn out that not all members of the P. parnellii group are high duty echolocators?

REFERENCES

de la Torre, J. A., & R. A. Medellín. 2010. Pteronotus personatus (Chiroptera: Mormoopidae). Mammalian Species 42 (869): 244–250.

Pavan, A. C., & G. Marroig. 2016. Integrating multiple evidences in taxonomy: species diversity and phylogeny of mustached bats (Mormoopidae: Pteronotus). Molecular Phylogenetics and Evolution 103: 184–198. Simmons, N. B., & T. M. Conway. 2001. Phylogenetic relationships of mormoopid bats (Chiroptera: Mormoopidae) based on morphological data. Bulletin of the American Museum of Natural History 258: 1–97.

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.