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Anomalurus beecrofti Fraser, 1852 Anomalurus beecrofti Fraser, 1852: 17. COMMON NAME. — Beecroft’s Anomalure; EWE: Kasanui, Vava. MATERIAL EXAMINED. — Agumatsa Wildlife Sanctuary • 1 ♀; ZTNHC 950. REMARK This specimen was brought to us by a local hunter, on 19 November 1999. In the Volta Region a previous record is from the vicinity of Jasikan (Grubb et al. 1998) and it was observed at Kyabobo National Park by Hurst et al. (1995). In the Togo Highlands A. beecrofti was previously recorded from Apéyémé, Edifou, Ezimé and Misahohé (Robbins & Van der Straeten 1996; USNM). Grubb et al. (1998) followed Rosevear (1969) in using the generic name Anomalurops Matschie, 1914 for this species; we follow Dieterlen (2005) and Kingdon (2013) in the use of Anomalurus Waterhouse, 1843. CONSERVATION STATUS. — Anomalurus beecrofti is listed as “Least Concern” on the IUCN Red List, however, progressive forest fragmentation with the loss of mature trees, specifically oil palms (Elaeis guineensis), will greatly compromise the habitat of this treedependent glider., Published as part of Decher, Jan, Norris, Ryan W., Abedi-Lartey, Michael, Oppong, James, Hutterer, Rainer, Weinbrenner, Martin, Koch, Martin, Podsiadlowski, Lars & Kilpatrick, C. William, 2021, A survey of small mammals in the Volta Region of Ghana with comments on zoogeography and conservation, pp. 253-281 in Zoosystema 43 (14) on page 265, DOI: 10.5252/zoosystema2021v43a14, http://zenodo.org/record/4783781, {"references":["FRASER L. 1852. - Description of a new species on Anomalurus from Fernando Po. Proceedings of the Zoological Society of London 20: 16 - 17. https: // www. biodiversitylibrary. org / page / 30680608","GRUBB P., JONES T. S., DAVIES A. G., EDBERG E., STARIN E. D. & HILL J. E. 1998. - Mammals of Ghana, Sierra Leone and the Gambia. The Trendrine Press, Zennor, St. Ives, Cornwall, vi + 265 p.","HURST J., BOWES LYON D. & KEEFE E. O' 1995. - Kyabobo' 94: a joint Ghanaian and Oxford University Expedition. Final Report. KCP Publications, Ghana Wildlife Department, Institute for Renewable Natural Resources, 69 p.","ROBBINS C. B. & VAN DER STRAETEN E. 1996. - Small mammals of Togo and Benin. II. Rodentia. Mammalia 60: 231 - 242.","ROSEVEAR D. R. 1969. - The rodents of West Africa. Trustees of the British Museum of Natural History, London, xii + 604 p.","DIETERLEN F. 2005. - Family Anomaluridae in WILSON D. E. & REEDEr D. M. (eds) Mammal species of the world. Johns Hopkins University Press, Baltimore: 1532 - 1534."]}

Five of the seven modern species of the family Anomaluridae have been studied. Significant intergeneric differences in the structure of the tongue between Anomalurus and Idiurus have been identified. These differences affect even such taxonomically important features as the number of circumvallate papillae (three in Anomalurus and two in Idiurus) and the distribution pattern of fungiform papillae. A parallel interspecific variability has been observed within the genera. It manifests itself in a decrease in the total number of fungiform papillae. The tongue structure in Anomalurus corresponds to the generalized version, initial for the order, whereas in Idiurus, it has clear signs of morphological and functional specializations.

A more anteriorly positioned foramen magnum evolved in concert with bipedalism at least four times within Mammalia: once in macropodid marsupials, once in heteromyid rodents, once in dipodid rodents, and once in hominoid primates. Here, we expand upon previous research on the factors influencing mammalian foramen magnum position (FMP) and angle with four new analyses. First, we quantify FMP using a metric (basioccipital ratio) not previously examined in a broad comparative sample of mammals. Second, we evaluate the potential influence of relative brain size on both FMP and foramen magnum angle (FMA). Third, we assess FMP in an additional rodent clade (Anomaluroidea) containing bipedal springhares (Pedetes spp.) and gliding/quadrupedal anomalures (Anomalurus spp.). Fourth, we determine the relationship between measures of FMP and FMA in extant hominoids and an expanded mammalian sample. Our results indicate that bipedal/orthograde mammals have shorter basioccipitals than their quadrupedal/non-orthograde relatives. Brain size alone has no discernible effect on FMP or FMA. Brain size relative to palate size has a weak influence on FMP in some clades, but effects are not evident in all metrics of FMP and are inconsistent among clades. Among anomaluroids, bipedal Pedetes exhibits a more anterior FMP than gliding/quadrupedal Anomalurus. The relationship between FMA and FMP in hominoids depends on the metric chosen for quantifying FMP, and if modern humans are included in the sample. However, the relationship between FMA and FMP is nonexistent or weak across rodents, marsupials, and, to a lesser extent, strepsirrhine primates. These results provide further evidence that bipedal mammals tend to have more anteriorly positioned foramina magna than their quadrupedal close relatives. Our findings also suggest that the evolution of FMP and FMA in hominins may not be closely coupled.

International audience; Anomaluroidea, commonly known as the “scaly-tailed squirrels,” are an emblematic group of tropical African mammals that includes gliding forms. The family Anomaluridae was until recently represented by three genera: the flying scaly-tailed squirrels (Anomalurus), the flying mouse (Idiurus) and the flightless scaly-tailed squirrels (Zenkerella). Idiurus and Zenkerella have long been grouped into the Zenkerellinae subfamily, and Zenkerella was interpreted as a rare case of evolutionary reversal to non-gliding lifestyle. Recent studies have demonstrated that Zenkerella is sister to all other modern anomalurids, and represents in fact the monogeneric family Zenkerellidae. The Anomalurus genus was split into Anomalurus and Anomalurops, but no study has ever considered all Anomalurus species together in a phylogeny to test the status of Anomalurops. Here, we used mitogenomic next-generation sequencing to infer the phylogenetic relationships among all extant anomalurids and to estimate their divergence ages. We found that the arboreal Zenkerella is the sister group of all extant gliding anomalurids (Idiurus and Anomalurus). We confirmed that Anomaluroidea only evolved the gliding adaptation once. A comparison based on morphological traits indicates that Zenkerella harbours several unique morphological features. We propose new morphological characters for the novel classification of modern Anomaluroidea, which includes the families Zenkerellidae and Anomaluridae. Using different calibration schemes, we demonstrated that classical dating methods relying only on mitogenomes provide rather young Miocene estimates between Zenkerellidae and the Anomaluridae. The use of published nuclear genes, internal calibrations and tip dating converged towards an Eocene split between gliding and non-gliding scaly-tailed squirrels, which is in agreement with the African fossil record. Finally, we provide the first exhaustive species-level molecular phylogenetic inference for the genus Anomalurus. We found that Anomalurus beecrofti is the sister group of all other species of Anomalurus and branched off during the Miocene.

The “scaly-tailed squirrels” of the rodent family Anomaluridae have a long evolutionary history in Africa, and are now represented by two gliding genera (AnomalurusandIdiurus) and a rare and obscure genus (Zenkerella) that has never been observed alive by mammalogists.Zenkerellashows no anatomical adaptations for gliding, but has traditionally been grouped with the gliderIdiuruson the basis of craniodental similarities, implying that either theZenkerellalineage lost its gliding adaptations, or thatAnomalurusandIdiurusevolved theirs independently. Here we present the first nuclear and mitochondrial DNA sequences ofZenkerella, based on recently recovered whole-body specimens from Bioko Island (Equatorial Guinea), which show unambiguously thatZenkerellais the sister taxon ofAnomalurusandIdiurus. These data indicate that gliding likely evolved only once within Anomaluridae, and that there were no subsequent evolutionary reversals. We combine this new molecular evidence with morphological data from living and extinct anomaluromorph rodents and estimate that the lineage leading toZenkerellahas been evolving independently in Africa since the early Eocene, approximately 49 million years ago. Recently discovered fossils further attest to the antiquity of the lineage leading toZenkerella, which can now be recognized as a classic example of a “living fossil,” about which we know remarkably little. The osteological markers of gliding are estimated to have evolved along the stem lineage of theAnomalurus–Idiurusclade by the early Oligocene, potentially indicating that this adaptation evolved in response to climatic perturbations at the Eocene–Oligocene boundary (∼34 million years ago).

3. Pel’s Anomalure Anomalurus pelii French: Anomalure de Pel / German: Pel-Dornschwanzhérnchen / Spanish: Anomaluro de Pel Other common names: Pel’s Flying Squirrel, Pel’s Scaly-tailed Flying Squirrel, Pel’s Scaly-tailed Squirrel Taxonomy. Pteromys pelii Schlegel & S. Miiller, 1845, “Daboerom, aan de Goudkust” (= Dabacrom, Ghana, West Africa). Three subspecies are recognized. Subspecies and Distribution. A.p.peliiSchlegel&Muller,1845—EIvoryCoast(EofBandamaRiver)andGhana(WofVoltaRiver). A.p.auzembergeriMatschie,1914—ELiberiaandextremeWIvoryCoast(WoftheSassandraRiver). A. p. peralbus Schunke & Hutterer, 2005 — Ivory Coast between Sassandra and Bandama rivers. Descriptive notes. Head-body 400-460 mm, tail 320-450 mm; weight 1.3-1.8 kg. Pel’s Anomalure is the largest species of anomalures and is immediately identified by its distinctive black color on dorsal surface, with varying degrees of white. The nominate subspecies pelii has a narrow white border on dorsal surface ofits patagium and a narrow border on the interfemoral membrane. Tail and hindfeet are whitish, forefeet are blackish, and there is a white patch at base of each ear. Patagium of the subspecies auzembergeri is black on dorsal surface without a white border, interfemoral membrane is black without a white border, and tail is black; there is no white patch at base of each ear and forefeet and hindfeet are black. Dorsal surface of patagium of the subspecies peralbus has a very wide white border that almost reaches the body; there is a wide white border on the interfemoral membrane;tail is white; there is a large white patch at base of each ear; and forefeet and hindfeet are white. Habitat. Moist, low-elevation primary forests, with numerous tall emergents and annual rainfall of 1400-3900 mm. Food and Feeding. Diet of Pel’s Anomalure consists of bark, supplemented by fruits, especially from African oil palm (Elaeis guineensis, Arecaceae), leaves, and flowers. Breeding. Pel’s Anomalures have two breeding seasons in April and September and produce 1-4 young/litter, more often 2-3. Activity patterns. Pel’s Anomalures are nocturnal and spend the day asleep in tree hollows or hollow branches. Nests include sticks and small branches. Sharing of dens by up to six individuals suggests that abundance might be strongly influenced by local availability oftree holes. One Pel’s Anomalure was found a hollow of Terminalia superba (Combretaceae). Pel’s Anomalures emerge from their dens well after sunset and return to them before sunrise. Movements, Home range and Social organization. Primary social organization of Pel’s Anomalure consists of a female and her single offspring, but there are records suggesting that up to six individuals can be found together in a sleeping tree hollow. When disturbed in their den, they are known to make several noises including growls and hisses; they snap their teeth, greatly amplified in the tree hollow. Contact calls of Pel’s Anomalures are described as deep hoots. These sounds may deter predators including birds of prey, primates, small carnivores, and hornbills (Bucerotidae) that may compete for nest holes. Bold color of Pel’s Anomalure suggests that aggressiveness and large size may make it less vulnerable to predation than smaller species of anomalures. Likely predators include African harrier-hawks (Polyboroides typus), crowned eagles (Stephanoaetus coronatus), hornbills, snakes, and African Palm Civets (Nandinia binotata). 2 Status and Conservation. Classified as Data Deficient on The IUCN Red List. It is likely that most populations of Pel’s Anomalures are threatened by the rapid rate of forest degradation in West Africa; surveys are needed to assess their status. To the Mbuti Pygmies in north-eastern DR Congo, skin of species of Anomalurus is burned and ground into a powder to make the drug “Bau,” which is blown into the face ofvictims at night, who then become unable to defend themselves. Bibliography. Adams (1894), Cansdale (1946), Carpaneto & Germi (1989), Corbet & Hill (1991), Dieterlen (1993), Dorst & Dandelot (1970), Grubb et al. (1998), Haltenorth & Diller (1977), Honacki et al. (1982), Jackson (2012), Jackson & Thorington (2012), Kingdon (1997), Rosevear (1969), Schunke (2005), Schunke & Hutterer (2004b, 2005)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Anomaluridae, pp. 270-279 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 278, DOI: 10.5281/zenodo.6584330

4. Lesser Anomalure Anomalurus pusillus French: Anomalure pygmée / German: Zwerg-Dornschwanzhérnchen / Spanish: Anomaluro enano Other common names: Dwarf Anomalure, Dwarf Scaly-tailed Flying Squirrel, Dwarf Scaly-tailed Squirrel, Little Flying Squirrel Taxonomy. Anomalurus pusillus Thomas, 1887, “Bellima and Tingasi, Monbuttu” (= Niangara, DR Congo). This species is monotypic. Distribution. Found in isolated populations in Liberia, Cameroon, Equatorial Guinea, NW Gabon, W Republic of the Congo, SW Central African Republic, and N DR Congo; may also occur in W Uganda. Descriptive notes. Head-body 190-260 mm, tail 120-200 mm; weight 170-300 g. The Lesser Anomalure is the smallest species of Anomalurus, with dorsal color varying from near black, grizzled gray, to mottled tan or olive brown. Membrane adjoining tail is usually yellower, and lateral membranes are dark gray. Head of the Lesser Anomalure is gray without borders around ears. Undersideis pale yellowish white, without a tinge of rufus; bristles covering hind claws have a rufus tinge. Unlike other species of Anomalurus, the Lesser Anomalure is similar in appearance throughoutits distribution. Habitat. Limited information available but only found in rainforest in Gabon and DR Congo. Most records of Lesser Anomalures have been at low elevations, but there is one record from montane forests at 2200 m. Food and Feeding. Diet of the Lesser Anomalure appears to include bark and fruits. Drupes of parasol trees (Musanga, Urticaceae) are eaten. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. Lesser Anomalures appear to nest alone. Density is only ¢.5 ind/km?. Status and Conservation. Classified as Least Concern on The IUCN Red List. Major threats to the Lesser Anomalure include loss of habitat as a result of conversion to agricultural and logging for timber. In some regions, they are hunted for bushmeat, but this is not thought to be a significant conservation threat. Bibliography. Delany (1975), Dorst & Dandelot (1970), Grubb et al. (1998), Haltenorth & Diller (1977), Jackson (2012), Jackson & Thorington (2012), Julliot et al. (1998), Kingdon (1997 2013d), Rahm (1969), Rahm & Christiaensen (1963), Schunke (2005)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Anomaluridae, pp. 270-279 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 278, DOI: 10.5281/zenodo.6584330

2. Lord Derby’s Anomalure Anomalurus derbianus French: Anomalure de Derby / German: Lord-Derby-Dornschwanzhérnchen / Spanish: Anomaluro de Lord Derby Other common names: Derby's Flying Squirrel, Lord Derby's Flying Squirrel, Lord Derby's Scaly-tailed Flying Squirrel, Lord Derby's Scaly-tailed Squirrel Taxonomy. Pteromys derbianus Gray, 1842, “W. Africa, Sierra Leone.” Although numerous forms have been attributed to A. derbianus, they are junior synonyms, and no subspecies are recognized. Monotypic. Distribution. W, C & E Africa, from Sierra Leone E to Uganda and SW Kenya, and S to Tanzania, N Mozambique, N Malawi, Zambia, and N Angola, also on Bioko I. Not yet recorded in Benin where may occur and possibly on Zanzibar I. Descriptive notes. Head-body 260-400 mm, tail 220-330 mm; weight 0-45—1-1 kg. Lord Derby’s Anomalure shows the highest variability in color in the family; it varies geographically and within populations. Lord Derby’s Anomalures range in color from more or less uniformly brownish to those with reddish backs, dark gray patagia, black ears, and silverfish stripes on nose. Head typically has a black-to-brown mask around eyes, muzzle, and surrounding the ears. Gliding membranes, which stretch between limbs and onto base of tail, are similarly colored above but black bristles reinforce edge of membrane behind elbow strut. Ventral surface is much less variable and patagium is generally white, but sometimes underside of membrane between hindlegs is brownish to blackish. Base oftail has 12-14 scales. Primary variation among Lord Derby’s Anomaluresis in gray or russet tints on back and shoulders. Face is relatively short and round; naked pinkish ears are set in a surrounding bed of dense velvet-black fur. Bristles covering hind claws are black. Habitat. Variety of forest types from moist rainforests to relatively dry savanna woodlands from sea level up to elevations of ¢.2400 m. In the south-eastern part of its wide distribution, Lord Derby’s Anomalure is known to occur in savanna woodland. Although they can be common in undisturbed forest, they may also nest within dead trees in areas that have been cleared and burned for agriculture, as long as trees are within gliding distance to nearby forest. Food and Feeding. Bark of trunk and main branches of several genera of trees appear to be the dietary staple of Lord Derby’s Anomalure. Feeding individuals appear to focus on bark wounds caused by falling branches, elephanttusks, and growth splits. As a consequence of this feeding behavior, some trees such as beli (Julbernardia pellegriniana, Fabaceae) may lose all their bark on their trunks and main branches over a number of years. Loss of bark appears to be tolerated by the tree because they are able rapidly to regrow their cambium layer, suggesting that these rodents and trees have coevolved. Lord Derby’s Anomalures eat fruits of Carapa (Meliaceae) and Syzygium (Myrtaceae), leaves of Cynometra (Fabaceae) and Myrianthus (Moraceae), and bark of various trees, including those of the genera Neoboutonia (Euphorbiaceae) Cynometra, Pentaclethra (Fabaceae), Klainedoxa (Irvingiaceae), Carapa, Syzygium, and Strombosia (Olacaceae) and Strombosia (Olacaceae). Elsewhere, they eat fruits, flowers, leaves, nuts, and occasional insects such as ants and termites. Breeding. Breeding of Lord Derby’s Anomalures appears to be seasonal in drier areas but not in the main forest block. In some areas, females are known to segregate to give birth and rear young. Pregnant females have been taken in Cameroon in June-July when rains begin, so young are probably born toward the end of rains in September— November. Other observations in Tanzania found well-grown young in May-June during the early dry season. In Equatorial Guinea, lactating females have been collected in February-May, and in Uganda and Kenya, four births were recorded in January, two pregnancies in March, and a lactating female in May. Vaginas of pregnant females are firmly sealed with hard plugs after mating. Generally only one young is born at a time, although there are records of 2-3 young. Offspring remain on a ledge high up in a hollow tree until almost fully grown. Both parents bring offspring well-masticated food, carried in their mouth and cheeks—swollen by a wad as large as a tangerine. Activity patterns. Lord Derby’s Anomalure is nocturnal, with individuals resting in hollow trees during the day, preferably in virgin forest. These hollows can be up to 40 m above the ground (in some woodlands they occupy empty beehives). One radiocollared individual in Gabon had one primary nesting tree hollow and five secondary hollows. They tend to nest alone or in pairs of one female and a young. Other observations suggest that up to eight individuals may share a denning tree, but little is known about their social behavior. Lord Derby’s Anomalures also appear to be sedentary, living in the same hole all year, except when fruit is ripe in another area. Individuals retain use of their dens over very long periods, but seasonal shifts have been recorded. Along with a threatening or defensive hiss and growl, Lord Derby’s Anomalures also purr and twitter. They like to sunbathe in early morning and more rarely in the evening. Pruning in closed-canopy forest is often less frequent or conspicuous than in more open or sunlit areas. Where light breaks through close to a food tree, intense pruning of young trees may help maintain access to a ready food supply by killing or maiming competing trees and keeping flight-paths open. Night is spent moving in lower branches oftall trees and smaller growth, often not more than 9 m above the ground. While resting, a Lord Derby’s Anomalure orientatesitself with head upward and body parallel with the bole of the tree. Movements, Home range and Social organization. When leaping from one tree to another, membranes of Lord Derby's Anomalures are curved such that an individual assumes shape of a small umbrella. They prefer to land on trunks rather than branches of trees. Immediately before reaching a tree trunk, head is actually at a lower level than tail, but at the last second, forelegs are thrown back over shoulders, head comes up, and tail sweeps up to meet it over the back. An individual instantly assumes a vertical position in midair. Momentum carries it to an upright trunk, to which it instantly adheres, and it invariably commences an ascent using its front two feet together and then pulling up the two hindfeet and arching its back like a giant loop-caterpillar. Backward-directed scales at base of the tail and forwardly directed claws on hindfeet dig into the three bark, thus forming a rigid tripod to support the individual while forefeet are released and moved upward. Glides of more than 40 m by Lord Derby’s Anomalures have been recorded at an estimated glide angle of 24° from horizontal. Species of Anomalurus can gallop up the smooth bark of a giant forest tree with great speed. Distance traveled per night averages 540 m; home range averages 3-4 ha for females and 3-3 ha for males. A radio-collared female had a home range of 3-4 ha; 50% of locations in a small central area comprised 11:3% of her home range. Densities of Lord Derby’s Anomalures are 27-54 ind/km?. Blue Monkeys (Cercopithecus mitis) prey on Lord Derby’s Anomalures in Budongo Forest Reserve, Uganda. Anomalure (Zenkerella insignis). Status and Conservation. Classified as Least Concern on The IUCN Red List. Although localized populations of Lord Derby’s Anomalures may be at risk from logging and forest clearance,it is common and widespread. It is also used in traditional medicine. Bibliography. Allen (1922), Delany (1975), Dorst & Dandelot (1970), Fairgrieve (1997), Grubb et al. (1998), Haltenorth & Diller (1977), Jackson (2012), Jackson & Thorington (2012), Jones (1971), Julliot et al. (1998), Kingdon (1974, 1997), Rahm (1969), Sanderson (1940), Schunke (2005)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Anomaluridae, pp. 270-279 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on pages 277-278, DOI: 10.5281/zenodo.6584330

1. Beecroft’s Anomalure Anomalurus beecrofty French: Anomalure de Beecroft / German: Beecroft-Dornschwanzérnchen / Spanish: Anomaluro de Beecroft Other common names: Beecroft's Flying Squirrel, Beecroft's Scaly-tailed Flying Squirrel, Beecroft's Scaly-tailed Squirrel Taxonomy. Anomalurus beecrofti Fraser, 1853, “Fernando Po” (Bioko Island, Equatorial Guinea, West Africa). Some authors have placed A. beecroftiin the genus Anomalurops, named by P. Matschie in 1914, by itself due to various skull characteristics. Three subspecies have been recognized including beecrofti from Bioko Island, Equatorial Guinea, named by L. Fraser in 1853; argenteus from Senegal to Cameroon, named by H. Schwann in 1904; and fulgens from Gabon, named by J. E. Gray in 1869. These subspecies typically are not recognized in recent publications. Monotypic. Distribution. Widely distributed in W & C Africa, from Senegal to Gabon, E DR Congo and Bioko I, also isolated records in NW Angola, NW Zambia, and W Uganda. Descriptive notes. Head—body 250-310 mm, tail 160-260 mm; weight 640-660 g. Beecroft’s Anomalureis large and similar in appearance to the smaller Lesser Anomalure (A. pusillus), which has relatively uniform dorsal fur and black-tipped tail. Lord Derby’s Anomalure (A. derbianus) is also similar to Beecroft’s Anomalure, but it has black or brown eye mask, muzzle, and area surrounding ears. Dorsal pelage of Beecroft’s Anomalure is thick, fleecy, and lightly waved; fur at base of tail is smooth-haired, with a slight terminal tuft. Ventral region and underside of gliding membrane are well furred. Color varies considerably but most commonly is silver-gray on back, with a central stripe of golden brown from neck over larger part of back. Ventral side is basically grayish or yellowish; throat and chest are orange. Color variation can be defined by extent of golden brown stripe on back and amount of orange on ventral side. Beecroft’s Anomalure often has a white spot on its forehead but does not have a dark mask around eyes. Tail is narrower than in other species of anomalures and has up to nine pairs of scales on its underside at its base. Habitat. Tropical moist forest and dry forests, preferring swamp forests and areas with abundant palms. Beecroft’s Anomalures are also commonly found in disturbed forest such as cultivated groves of African oil palm (Elaeis guineensis, Arecaceae). They occur from sea level to elevations of 2500 m. Food and Feeding. Beecroft’s Anomalures eat leaves, bark, and fruit from a variety of trees and occasionally insects. They feed on pulp offruit of the African oil palm, which they manage to access despite the protection of short spines. Breeding. Young Beecroft’s Anomalures are born singly and are carried by their mothers when they are small. There appears to be a prolonged breeding season at the end of the wet season. One pregnant female was captured in July in Cameroon; two females, each with one young, were obtained in early October and late January. Activity patterns. Beecroft’s Anomalures are nocturnal, and during the day, they rest in tree holes located in upper strata or hide among leaves and twigs that form dense congregations. They may also rest in junctions between palm fronds, or cling to sheltered undersides of major tree branches close to the trunk, where their mottled coat helps to camouflage them away from predators. A jaw fragment of Beecroft’s Anomalures was found in a Chimpanzee (Pan troglodytes) fecal sample in the Tenkere area in Sierra Leone. Other likely predators include crowned eagles (Stephanoaetus coronatus), long-tailed hawks (Urotriorchis macrourus), and pythons (Pythonidae). Movements, Home range and Social organization. Beecroft’s Anomalures are typically found alone, which suggest they are solitary. The only record of a vocalization is something between a hoot and a whistle. Status and Conservation. Classified as Least Concern on The IUCN Red List. Although Beecroft’s Anomalures are difficult to observe, they appear be relatively common in suitable habitats but may be declining locally in some areas. Bibliography. Allen, G.M. & Coolidge (1930), Allen, J.A. (1922), Alp & Kitchener (1993), Bates (1905), Dekeyser & Villiers (1951), Delany (1975), Dorst & Dandelot (1970), Fraser (1853), Gray (1869), Haltenorth & Diller (1977), Happold (1987), Jackson (2012), Jackson &Thorington (2012), Kingdon (1997, 2013b), Matschie (1914), Rosevear (1969), Sanderson (1940), Schunke (2005), Schwann (1904)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Anomaluridae, pp. 270-279 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 277, DOI: 10.5281/zenodo.6584330

A new genus and species of diminutive anomalurid rodent, Shazurus minutus, is described on the basis of 15 isolated teeth from the earliest late Eocene (approximately 37 Ma) Birket Qarun Locality 2 in the Fayum Depression of northern Egypt. Shazurus is surprisingly specialized for its age, being most similar in dental morphology to early Miocene Paranomalurus and extant Anomalurus, and is quite different from the roughly contemporaneous anomaluroid genera Nementchamys and Pondaungimys from Algeria and Myanmar, respectively. Parsimony analysis of dental features places Shazurus within crown Anomaluridae as a sister group of two species of Paranomalurus to the exclusion of extant Anomalurus and Idiurus. The marked morphological differences between the two oldest Afro-Arabian anomaluroids (Shazurus and Nementchamys) suggest that the taxa share a much more ancient common Afro-Arabian ancestor, possibly derived from Zegdoumyidae. Isolated teeth of Shazurus and other Eocene anomaluroids reveal little about their palaeobiology, but the complete absence of Anomaluroidea from the younger (late Eocene to early Oligocene) Jebel Qatrani Formation is presumably a reflection of environmental change through the late Eocene in northern Africa.

Anomaluroid rodents show interesting biogeographic and macroevolutionary patterns, although their fossil record is meager and knowledge of the natural history of extant members of the clade remains inadequate. Living anomaluroids (Anomaluridae) are confined to equatorial parts of western and central Africa, but the oldest known fossil anomaluroid (Pondaungimys) comes from the late middle Eocene of Myanmar. The first appearance of anomaluroids in the African fossil record coincides with the first appearances of hystricognathous rodents and anthropoid primates there. Both of the latter taxa are widely acknowledged to have originated in Asia, suggesting that anomaluroids may show a concordant biogeographic pattern. Here we describe two new taxa of African Paleogene anomaluroids from sites in the Sirt Basin of central Libya. These include a new Eocene species of the nementchamyid genus Kabirmys, which ranks among the oldest African anomaluroids recovered to date, and a new genus and species of Anomaluridae from the early Oligocene, which appears to be closely related to extant Zenkerella, the only living non-volant anomalurid. Phylogenetic analyses incorporating the new Libyan fossils suggest that anomaluroids are not specially related to Zegdoumyidae, which are the only African rodents known to antedate the first appearance of anomaluroids there. The evolution of gliding locomotion in Anomaluridae appears to conflict with traditional assessments of relationships among living anomalurid taxa. If the historically accepted division of Anomaluridae into Anomalurinae (extant and Miocene Anomalurus and Miocene Paranomalurus) and Zenkerellinae (extant and Miocene Zenkerella and extant Idiurus) is correct, then either gliding locomotion evolved independently in Anomalurinae and Idiurus or non-volant Zenkerella evolved from a gliding ancestor. Anatomical data related to gliding in Anomaluridae are more consistent with a non-traditional systematic arrangement, whereby non-volant Zenkerella is the sister group of a clade including both Anomalurus and Idiurus. Kabirmys prius: urn:lsid:zoobank.org:act:C4FC0114-DAA6-4D12-BD3C-F5900BE1AD1E Prozenkerella: urn:lsid:zoobank.org:act:144DC508-333F-4746-A313-A0556804D7F1 Prozenkerella saharaensis: urn:lsid:zoobank.org:act:76992C9D-D754-40E9-B8E2-E786A5BC7E8C urn:lsid:zoobank.org:pub:866B8A65-BCB2-4E8F-B6FA-24637606528C

An RNA-Seq strategy was used to obtain the complete set of protein-coding mitochondrial genes from two rodent taxa. Thanks to the next generation sequencing (NGS) 454 approach, we determined the complete mitochondrial DNA genome from Graphiurus kelleni (Mammalia: Rodentia: Gliridae) and partial mitogenome from Pedetes capensis (Pedetidae), and compared them with published rodent and outgroup mitogenomes. We finished the mitogenome sequencing by a series of amplicons using conserved PCR primers to fill the gaps corresponding to tRNA, rRNA and control regions. Phylogenetic analyses of the mitogenomes suggest a well-supported rodent phylogeny in agreement with nuclear gene trees. Pedetes groups with Anomalurus into the clade Anomaluromorpha, while Graphiurus branches within the squirrel-related clade. Moreover, Pedetes+Anomalurus branch with Castor into the mouse-related clade. Our study demonstrates the utility of NGS for obtaining new mitochondrial genomes as well as the importance of choosing adequate models of sequence evolution to infer the phylogeny of rodents.

Preliminary excavations in western Kenya from the middle Miocene Muruyur Beds, in particular from the Kipsaramon site complex (15.5 Ma), are yielding a diverse rodent fauna from a poorly documented period of African prehistory. Seven families (?sciurids, pedetids, anomalurids, thryonomyids, diamantomyids, myophiomyids, and cricetodontids) are represented. Thryonomyoids are dominant; cricetids and murids of modern aspect are currently unknown. The fauna includes a mixture of primitive early Miocene taxa (e.g., Diamantomys and Notocricetodon) and also more derived taxa (i.e., Thryonomyidae genus and species large). A new small anomalurid, Anomalurus parvus, sp. nov., is reported. Faunal composition and the distribution of rodent taxa from the Muruyur Beds show similarities with early and middle Miocene sites in East Africa, Namibia, and Saudi Arabia. Comparison of early and middle Miocene African rodent faunas suggests a period of relative faunal stability among rodent taxa during this time.

Anomalurus beecrofti Fraser, 1853. Proc. Zool. Soc. Lond., 1852:17 [1853]. TYPE LOCALITY: Equatorial Guinea, "Fernando Po" (= Bioko). DISTRIBUTION: Senegal to Uganda and Zaire (Kasai); Bioko (Equatorial Guinea). STATUS: CITES - Appendix III (Ghana). SYNONYMS: argenteus Schwann, 1904; chapini J. A. Allen, 1922; citrinus Thomas, 1916; fulgens Gray, 1869; hervoi Dekeyser and Villiers, 1951; laniger Temminck, 1853; schoutedeni Verheyen, 1968. COMMENTS: See Misonne (1974:4). Matschie (1914) placed beecrofti in his subgenus Anomalurops. Includes schoutedeni which is considered a distinct species by Cabral (1971)., Published as part of Fritz Dieterlen, 1993, Order Rodentia - Family Anomaluridae, pp. 757-758 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 757, DOI: 10.5281/zenodo.7359185, {"references":["Gray, J. E. 1869. Catalogue of carnivorous, pachydermatous and edentate mammals in the British Museum. British Museum (Natural History) Publications, London, 398 pp.","Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated.","Matschie, P. 1914. Ein neuer Anomalurus von der Elfenbeinkuste. Sitzungs-berichte der Gesellschaft Naturforschender Freunde zu Berlin, 1914: 349 - 351.","Cabral, J. C. 1971. Existencia em Angola de Anomalurops beecrofti (Fraser). Boletim do Instituto de Investigacao Cientifica de Angola, 8: 55 - 63."]}

Anomalurus derbianus (Gray, 1842). Ann. Mag. Nat. Hist., [ser. 1], 10:262. TYPE LOCALITY: " Sierra Leone ". DISTRIBUTION: Sierra Leone to Angola, east to Kenya, south to Mozambique and Zambia. STATUS: CITES - Appendix III (Ghana). SYNONYMS: beldeni Du Chaillu, 1860; chrysophaenus Dubois, 1888; cinereus Thomas, 1895; erythronotus Milne-Edwards, 1879; fortior Lönnberg, 1917; fraseri Waterhouse, 1842; griselda Dollman, 1914; imperator Dollman, 1911; jacksoni de Winton, 1898; jordani St. Leger, 1935; laticeps Aguilar-Amat, 1922; neavei Dollman, 1909; nigrensis Thomas, 1904; orientalis Peters, 1880; perustus Thomas, 1914; squamicaudus Schinz, 1845. COMMENTS: See Misonne (1974:4-5)., Published as part of Fritz Dieterlen, 1993, Order Rodentia - Family Anomaluridae, pp. 757-758 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 757, DOI: 10.5281/zenodo.7359185, {"references":["Thomas, O. 1895. Preliminary diagnoses of new mammals from northern Luzon, collected by Mr. John Whitehead. Annals and Magazine of Natural History, ser. 6, 16: 160 - 164.","Waterhouse, G. R. 1842 [1843]. Descriptions of a new species of quadrupeds collected by Mr. Fraser at Fernando Po. Proceedings of the Zoological Society of London, 1842: 124 - 130.","Dollman, G. 1911. On Arvicanthis abyssinicus and allied East-African species, with descriptions of four new forms. Annals and Magazine of Natural History, ser. 8, 8: 334 - 353.","Dollman, G. 1909. New mammals from British East Africa. Annals and Magazine of Natural History, ser. 8, 4: 39 - 553.","Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

Anomalurus pusillus Thomas, 1887. Ann. Mag. Nat. Hist., ser. 5, 20:440. TYPE LOCALITY: Zaire, "Beilima and Tingasi, Monbuttu". DISTRIBUTION: S Cameroon, Gabon, Zaire. SYNONYMS: batesi de Winton, 1897. COMMENTS: See Misonne (1974:5)., Published as part of Fritz Dieterlen, 1993, Order Rodentia - Family Anomaluridae, pp. 757-758 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 757, DOI: 10.5281/zenodo.7359185, {"references":["Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

Anomalurus pelii (Schlegel and Müller, 1845). In Temminck, Verh. Nat. Gesch. Nederland. Overz. Bezitt., Zool., Mammalia, p. 109 [1845]. TYPE LOCALITY: "bÿ Daboerom, aan de Goudkust" (= Ghana, Dabacrom). DISTRIBUTION: Sierra Leone to Ghana. STATUS: CITES - Appendix III (Ghana). SYNONYMS: auzembergeri Matschie, 1914. COMMENTS: See Misonne (1974:4)., Published as part of Fritz Dieterlen, 1993, Order Rodentia - Family Anomaluridae, pp. 757-758 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 757, DOI: 10.5281/zenodo.7359185, {"references":["Matschie, P. 1914. Ein neuer Anomalurus von der Elfenbeinkuste. Sitzungs-berichte der Gesellschaft Naturforschender Freunde zu Berlin, 1914: 349 - 351.","Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

Anomalurus Waterhouse, 1843. Proc. Zool. Soc. Lond., 1842: 124 [1843]. TYPE SPECIES: Anomalurus fraseri Waterhouse, 1843 (= Pteromys derbianus Gray, 1842). SYNONYMS: Anomalurella Matschie, 1914; Anomalurodon Matschie, 1914; Anomalurops Matschie, 1914; Aroaethrus Waterhouse, 1843. COMMENTS: See Misonne (1974:3-5)., Published as part of Fritz Dieterlen, 1993, Order Rodentia - Family Anomaluridae, pp. 757-758 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 757, DOI: 10.5281/zenodo.7359185, {"references":["Gray, J. E. 1842. Description of two new species of Mammalia discovered in Australia by Captain George Gray (Tarsipes Spenserae and Chaeropus castanotis). Annals and Magazine of Natural History, [ser. 1], 9: 39 - 42.","Matschie, P. 1914. Ein neuer Anomalurus von der Elfenbeinkuste. Sitzungs-berichte der Gesellschaft Naturforschender Freunde zu Berlin, 1914: 349 - 351.","Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

Unworn teeth are figured and described for Anomalurus , Petromus and Thryonomys , and partly worn teeth for Pedetes . Anomalurus has five-crested teeth, although the teeth of the related Anomalurops have only four. It is impossible to determine which is the primitive number, and hence the ancestry of the family is uncertain. Petromus has four-crested teeth. The upper teeth of Thryonomys are four-crested; the lower molars have only three crests. Both of these families seem to have had a four-crested ancestry, and probably are related at the superfamily level. The ctenodactylids also have a four-crested pattern, but otherwise do not seem close to the Thryonomyoidea. The tooth pattern of Pedetes is based on two transverse rows of two cusps each, unlike that of any other rodents. The pedetids are completely isolated.

Anomalurus Waterhouse, 1843. Proc. Zool. Soc. Lond., 1842:124. COMMENT: Includes Anomalurops; see Misonne, 1974, Part 6:4. PROTECTED STATUS: CITES - Appendix III as Anomalurus spp. native to Ghana only. ISIS NUMBER: 5301410006002000000 as Anomalurus. 5301410006001000000 as Anomalurops., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 2), pp. 382-392 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 390, DOI: 10.5281/zenodo.7353025, {"references":["Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

Anomalurus derbianus (Gray, 1842). Ann. Mag. Nat. Hist., ser. 1, 10: 262. TYPE LOCALITY: Sierra Leone. DISTRIBUTION: Sierra Leone to Angola, east to Kenya, south to Mozambique and Zambia. COMMENT: Includes cinereus, erythronotus, fraseri, imperator, jacksonii, neavei, and orientalis; see Misonne, 1974, Part 6:5. PROTECTED STATUS: CITES - Appendix III (Ghana). ISIS NUMBER: 5301410006002002001 as A. derbianus. 5301410006002001001 as A. cinereus. 5301410006002003001 as A. jacksoni (sic). 5301410006002004001 as A. orientalis., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 2), pp. 382-392 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 391, DOI: 10.5281/zenodo.7353025, {"references":["Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

Anomalurus beecrofti Fraser, 1853. Proc. Zool. Soc. Lond., 1852:17. TYPE LOCALITY: Equatorial Guinea, Bioko (= Fernando Po). DISTRIBUTION: Senegal to Uganda and Kasai (Zaire); Bioko. COMMENT: Includes fulgens; formerly included in Anomalurops; see Misonne, 1974, Part 6:4. PROTECTED STATUS: CITES - Appendix III (Ghana). ISIS NUMBER: 5301410006001001001 as Anomalurops beecrofti., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 2), pp. 382-392 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 391, DOI: 10.5281/zenodo.7353025, {"references":["Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

Anomalurus pusillus Thomas, 1887. Ann. Mag. Nat. Hist., ser. 5, 20:440. TYPE LOCALITY: Zaire, Monbuttu, Beilima and Tingasi. DISTRIBUTION: S. Cameroun; Gabon; Zaire. COMMENT: Includes batesi; see Misonne, 1974, Part 6:5. ISIS NUMBER: 5301410006002006001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 2), pp. 382-392 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 391, DOI: 10.5281/zenodo.7353025, {"references":["Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

Anomalurus peli (Temminck, 1845). Verhandl. Nat. Gesch. Nederland Bezitt., Zool., l (2):109. TYPE LOCALITY: Ghana (= Gold Coast), Dabocrom. DISTRIBUTION: Sierra Leone to Ghana. COMMENT: Includes auzembergeri; see Misonne, 1974, Part 6:4. PROTECTED STATUS: CITES - Appendix III (Ghana). ISIS NUMBER: 5301410006002005001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 2), pp. 382-392 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 391, DOI: 10.5281/zenodo.7353025, {"references":["Misonne, X. 1974. Order Rodentia. Part 6. Pp. 1 - 39, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.). [issued 10 Sep 1974]. Smithsonian Institution Press, Washington, D. C., not continuously paginated."]}

The affinities of the isolated rodent family Anomaluridae are unknown. Both sciuromorphous and specialized features characterize its placental morphology. A shallow preplacenta consists of inner cytotrophoblastic and outer syncytiotrophoblastic layers. The early chorio-allantoic placenta consists of the following zones: a hemochorial zone of interconnected trabeculae, lined with outer syncytiotrophoblastic and inner cytotrophoblastic layers; anchoring trabeculae attached to a zone of decidual giant cells, with the giant cells gradually disappearing in later stages; a zone of acidophilic, PAS-positive necrotic material underlying the giant cells; and a zone of decidua basalis underlying the necrotic zone and probably contributing to it. An allantoic vesicle lies on the surface of the early placenta; it is reduced in later stages. The bilaminar omphalopleure persists, and it is attached to the margins of the placental disc. The fetal membranes of Anomalurus are most similar to those of sciuromorphs, Pedetes and Ctenodactylus; this may be the result of primitive retentions.

The comparative myology of the gliding membranes of the marsupial genera Acrobates, Petauroides, Petaurus and Hemibelideus is compared with that of the skin musculature of the non-glider Pseudocheirus. An outline of the system used in the naming of the parts of the cutaneus maximus is included. The gliding membranes are compared, functionally, with those of the rodent Petauristinae; the positions of the muscles within the phalanger membranes account for the gliders' ability to execute changes in camber, thus lift and angle of attack. An accessory cartilage associated with the elbow of Petauroides helps to increase the size of the plagiopatagium; its anatomy is compared with that of a similar cartilage in the rodent genus Anomalurus. A scheme for the evolution of the gliders' dermal musculature from a Pseudocheirus- type ancestor is presented; it is suggested, because of its anatomy and habits, that Hemibelideus be elevated to full generic status.