Phylogeny and taxonomy of the round-eared sengis or elephant-shrews, genus Macroscelides (Mammalia, Afrotheria, Macroscelidea)

The round-eared sengis or elephant-shrews (genus Macroscelides) exhibit striking pelage variation throughout their ranges. Over ten taxonomic names have been proposed to describe this variation, but currently only two taxa are recognized (M. proboscideus proboscideus and M. p. flavicaudatus). Here, we review the taxonomic history of Macroscelides, and we use data on the geographic distribution, morphology, and mitochondrial DNA sequence to evaluate the current taxonomy. Our data support only two taxa that correspond to the currently recognized subspecies M. p. proboscideus and M. p. flavicaudatus. Mitochondrial haplotypes of these two taxa are reciprocally monophyletic with over 13% uncorrected sequence divergence between them. PCA analysis of 14 morphological characters (mostly cranial) grouped the two taxa into non-overlapping clusters, and body mass alone is a relatively reliable distinguishing character throughout much of Macroscelides range. Although fieldworkers were unable to find sympatric populations, the two taxa were found within 50 km of each other, and genetic analysis showed no evidence of gene flow. Based upon corroborating genetic data, morphological data, near sympatry with no evidence of gene flow, and differences in habitat use, we elevate these two forms to full species.     

New remains of Chambius kasserinensis from the Eocene of Tunisia and evaluation of proposed affinities for Macroscelidea (Mammalia,Afrotheria)

Abstract

Chambius kasserinensis from the late Early or early Middle Eocene Chambi locality, central Tunisia, is undoubtedly the oldest known macroscelidid and possibly the basalmost representative of the order Macroscelidea. Hence, since its discovery in 1986, Chambius has played a key role in analyses focusing on afrotherian and eutherian phylogeny; for instance, as early as 1995, Butler’s review of fossil macroscelideans highlighted the central position of Chambius in the origin of the order. Despite this, Chambius remained poorly known until recently. Here based on new mandibular fragments, well-preserved upper molars and CT scan analysis of the holotype maxilla, Chambius is revised. Its dentition is first described in detail, providing a precise characterization of the genus. Chambius is notably defined by a submolariform P₄ with a three-cusped talonid, a reduced talonid on M₂, and a prominent metaconule on M^1?2. Interestingly, the two transverse lophs of the upper molars are basically formed by preconulecristae, evoking the recently defined peculiar bilophodonty of paenungulates. Comparisons with other Paleogene and modern macroscelidids, European Louisinidae, and North American Apheliscidae are also made, allowing the various hypotheses about the origin and early evolution of macroscelidids to be reviewed.     

Confirmation of pleisiomorphic daily torpor in mammals: the round-eared elephant shrew Macroscelides proboscideus (Macroscelidea)

The characteristics of daily torpor were measured in the round-eared elephant shrew Macroscelides proboscideus (Macroscelidea) in response to ambient temperature and food deprivation. Elephant shrews are an ancient mammal order within a superordinal African clade including hyraxes, elephants, dugongs and the aardvark. M. proboscideus only employed torpor when deprived of food; torpor did not occur under an ad libitum diet at ambient temperatures of 10, 15 and 25?°C. Torpor bout duration ranged from <1?h to ca. 18?h. The times of entry into torpor were restricted to the scotophase, despite normothermic body temperature patterns indicating a rest phase coincident with the photophase. Full arousal was always achieved within the first 3?h of the photophase. When food deprived, the onset of the rest phase, and hence torpor, advanced with respect to the experimental photoperiod. The lowest torpor body temperature measured was 9.41?°C. Daily torpor in M. proboscideus confirms a pleisiomorphic origin of daily heterothermy. Torpor facilitates risk-averse foraging behaviour in these small omnivores by overcoming long-term energy shortfalls generated by the inherent variability of food availability in their semi-arid, El Niño-afflicted habitats.     

Morphological cranial diversity contributes to phylogeny in soft-furred sengis (Afrotheria, Macroscelidea)

Despite the well-supported Macroscelidea phylogeny proposed at the end of the 1960s, several systematic arrangements have been suggested in the last 20 years, raising doubts about the phylogeny of the Macroscelidinae; sengi inter-specific relationships are still debated to this day. The main issue of concern involves the supposed Elephantulus diphyly. To solve this persisting debate about sengi phylogeny, we examined the cranium ventral surface of 13 species using geometric morphometric techniques and neighbour-joining algorithms. This study supported the idea that the ventral side of the sengi cranium has the potential to provide important signals for reconstructing the Macroscelidea phylogeny. The phylogenetic signals seemed to differentiate between two major clades in the sengi radiation. In the first clade, the two monospecific genera (Petrodromus and Macroscelides), the two African Horn species (Elephantulus revoilii and E. rufescens), and the only North African species (E. rozeti) were clustered together. The second clade includes the remnant south-central African Elephantulus species. Our results were in agreement with both mitochondrial and nuclear data, confirmed that there is no Elephantulus monophyly and highlighted the close relationship between Petrodromus and E. rozeti. It appears that all the soft-furred sengi species are organised in two evolutionary lines: an old monophyletic clade, comprising only Elephantulus species, and a new polyphyletic clade, including P. tetradactylus, M. proboscideus, and E. rozeti. This requires a taxonomic and nomenclatural rearrangement within Macroscelidinae, where the phylogenetic position of the remnant 4 (of 12) Elephantulus species has yet to be fully defined.     

Why is there discordant diversity in sengi (Mammalia: Afrotheria: Macroscelidea) taxonomy and ecology?

The seventeen species of sengis or elephant-shrews form a well-defined clade of mammals endemic to Africa that occupy the extremes of terrestrial habitats, from coastal deserts to montane forests. Because of their isolation on Africa soon after the break-up of Gondwanaland, theoretically sengis initially evolved with little competition from other placental radiations. Their life history features include myrmecophagy, saltatorial gaits, no or limited use of nests, social monogamy, small litters of precocial young and absentee maternal care of neonates. These traits together are unique to the Macroscelidea and represent a wedding of features usually associated with either small antelopes or anteaters. Combined, these features define an adaptive syndrome that presumably has been relatively immune to competition from contemporary mammals, partially due to phylogenetic inertia. Yet paradoxically, the syndrome is well suited to a wide range of terrestrial habitats, resulting in low taxonomic diversity. Because of their unusual phylogeny and low species diversity, conservation interest is high for those sengis with relatively low densities in fragmented forests.     

New stem elephant‐shrews (Mammalia,Macroscelidea) from the Eocene of Dur At‐Talah,Libya

Abstract: Two new primitive macroscelideans, Eotmantsoius perseverans gen. et sp. nov. and Nementchatherium rathbuni sp. nov. from the late Middle or Late Eocene locality of Dur At‐Talah, Libya, are described. Eotmantsoius is known from a single tooth and is characterized by low crown height, isolated conules, and lack of a preprotocrista and prehypocrista. Based on these characters, it appears to be more plesiomorphic than Chambius (late Early Eocene, Chambi, Tunisia), which was, to date, universally accepted as the oldest and most basal macroscelidean. Eotmantsoius appears, however, somewhat derived in having an entostyle, a pericone, a shorter mesial cingulum and larger intercusp valleys. The phylogenetic position of Eotmantsoius is unclear. The second new species, N. rathbuni, allows a reassessment of Nementchatherium. We suggest that it is the most derived of the herodotiines in having long preprotocrista and prehypocrista, reduced M2/m2, a robust entostyle twinned with the hypocone on M1 and the posterior aspect of the root of the maxillary jugal process placed along the length of M2. Nementchatherium rathbuni appears more derived than N. senarhense, from the Eocene locality of Bir El Ater, Algeria, in having smaller m2‐3 with no paraconid, a smaller hypoconulid on m2 and a more reduced talonid on m3 (with smaller hypoconid and entoconid). Nementchatherium rathbuni also has, distal to the m3, a coronoid canal. This character has commonly been proposed as a synapomorphy for paenungulates (e.g. proboscideans, sirenians and hyracoids); we investigate the possibility that it unites macroscelideans and paenungulates, as the superordinal clade Afrotheria suggests. Comparisons reveal that a coronoid canal also occurs in several nonafrotherian orders (lagomorphs, xenarthrans, artiodactyls and perissodactyls), suggesting that is not an exclusive synapomorphy for paenungulates plus macroscelideans. Moreover, the homology of the character within paenungulates is doubtful.     

Daily torpor in elephant shrews (Macroscelidea: Elephantulus spp.) in response to food deprivation

Patterns of daily torpor were measured in response to photoperiod and food restriction at a constant temperature (18 °C) in two species of elephant shrew (Macroscelidea), Elephantulus rozeti (from Morocco) and Elephantulus myurus (from southern Africa). Body temperature was monitored continuously for ca. 3 months using temperature-sensitive telemeters. Under short photoperiods (8:16 L:D), both species entered spontaneous torpor on an ad libitum diet, but showed a higher frequency of induced torpor when food was restricted. Under long photoperiods (16:8 L:D), E. myurus could be induced to enter daily `summer' torpor. A total of 378 torpor bouts were measured, none of which were longer in duration than 18 h. Under short photoperiods, arousal from torpor was associated with the onset of the photoperiod, whereas the time of entry was variable throughout the scotophase. However, E. myurus tended to phase shift torpor from the photophase to the scotophase under long photoperiods, despite displaying weak circadian amplitudes of body temperature indicative of a photophase rest phase. Both species lacked well-defined circadian amplitudes of body temperature, a pattern thought to be associated with polyphasic activity cycles characteristic of several Elephantulus species. It was concluded that these and other patterns of torpor shown by Elephantulus show similarities with other small Afrotropical insectivores inhabiting semi-arid habitats or unpredictable environments. Accepted: 26 July 2000     

Shape and size variations in the cranium of elephant-shrews: a morphometric contribution to a phylogenetic debate

A geometric morphometric analysis was carried out on the crania of 13 species of elephant-shrews (Macroscelidea), a group of African mammals whose phylogeny is still debated. The material examined consisted of 313 crania and included all the genera of Macroscelididae, the unique family recognized by taxonomists. The results obtained from the analysis of the cranium shape and size, either from dorsal or lateral view, were very similar. The first one appeared more reliable because of the higher number of intersection points fixed between the cranial sutures. All the cranial features that distinguished the genus Rhynchocyon were a consequence of the extreme enlargement of frontal bones. Instead, within the subfamily Macroscelidinae, the differences between genera were based on modification involving other bones, mainly mastoids and nasals, as shown by the deformation grids. A cluster analysis confirmed the traditional subdivision in two subfamilies (Rhynchocyoninae and Macroscelidinae) but suggested a different relationship among the recognized genera belonging to Macroscelidinae. Our results are congruent with data obtained from previous biochemical research and support the traditional subdivision in two subfamilies, the monophily of the genus Elephantulus and its closeness with Petrodromus, relating to their similar cranium shapes. The latter presumably is a case of gigantism as adaptation to forest habitats. Further studies on all the species of Elephantulus could provide new evidence for assessing the relationships within this clade, including Macroscelides that by the present analysis appeared as a well-distinguished taxonomic entity.     

Nectar feeding by the Cape rock elephant-shrew Elephantulus edwardii (Macroscelidea) — A primarily insectivorous mammal pollinates the parasite Hyobanche atropurpurea (Orobanchaceae)

Cape rock elephant-shrews, Elephantulus edwardii (Macroscelidea) were found to be pollinators of Hyobanche atropurpurea (Orobanchaceae), a member of a holoparasitic plant genus, lacking up to now any detailed pollinator observations. The elephant-shrews were live-trapped near the plants in the Western Cape of South Africa. One animal was released in a terrarium, containing the plants. It licked nectar with its long and slender tongue while being dusted with pollen and touching the stigmas of the flowers with its nose and the fur above the snout. H. atropurpurea pollen was found in swabs of the fur around noses and snouts as well as in the faeces of captured animals. The occurrence of pollen in the faeces of the elephant-shrews is likely a result of grooming their fur as they visited the flowers without eating or damaging them. This is one of the first records of pollinating, nectar-drinking elephant-shrews, contributing to the very scarce knowledge about the behaviour of those primarily insectivorous animals, as well as small mammal pollination.     

Towards the bibliography of life

This paper discusses how we intend to take forward the vision of a Bibliography of Life in the ViBRANT project. The underlying principle of the Bibliography is to provide taxonomists and others with a freely accessible bibliography covering the whole of life. Such a bibliography has been achieved for specific study areas within taxonomy, but not for "life" as a whole.The creation of such a comprehensive tool has been hindered by various social and technical issues. The social concerns focus on the willingness of users to contribute to the Bibliography. The technical concerns relate to the architecture required to deliver the Bibliography. These issues are discussed in the paper and approaches to addressing them within the ViBRANT project are described, to demonstrate how we can now seriously consider building a Bibliography of Life. We are particularly interested in the potential of the resulting tool to improve the quality of bibliographic references. Through analysing the large number of references in the Bibliography we will be able to add metadata by resolving known issues such as geographical name variations. This should result in a tool that will assist taxonomists in two ways. Firstly, it will be easier for them to discover relevant literature, especially pre-digital literature; and secondly, it will be easier for them to identify the canonical form for a citationThe paper also covers related issues relevant to building the tool in ViBRANT, including implementation and copyright, with suggestions as to how we could address them.     

Edith R. Saunders and floral anatomy: bibliography and index to families she studied

Edith R. Saunders' contributions to floral anatomy are briefly considered. Fourteen of her abstracts and discussions and 49 of her lengthier works are cited. An index to 249 families of angiosperms sensu Airy Shaw (1973) keyed to Saunders' publications is given.     

A new species of giant sengi or elephant-shrew (genus Rhynchocyon) highlights the exceptional biodiversity of the Udzungwa Mountains of Tanzania

A new species of sengi, or elephant-shrew, is described. It was discovered in the northern Udzungwa Mountains of Tanzania in 2005. Sengis (Order Macroscelidea, super-cohort Afrotheria) include four genera and 15 species of mammals that are endemic to Africa. This discovery is a significant contribution to the systematics of this small order. Based on 49 camera trap images, 40 sightings and five voucher specimens, the new sengi is diurnal and distinguished from the other three species of Rhynchocyon by a grizzled grey face, pale yellow to cream chest and chin, orange-rufous sides, maroon back and jet-black lower rump and thighs. The body weight of the new species is about 700 g, which is 25–50% greater than any other giant sengi. The new Rhynchocyon is only known from two populations that cover about 300 km² of montane forest. It has an estimated density of 50–80 individuals km⁻². This discovery has important implications for the conservation of the high biodiversity that is found in the forests of the Eastern Arc Mountains.     

A catalogue of Lithuanian beetles (Insecta, Coleoptera)

This paper presents the first complete and updated list of all 3597 species of beetles (Insecta: Coleoptera) belonging to 92 familiesfound and published in Lithuania until 2011, with comments also provided on the main systematic and nomenclatural changes since the last monographic treatment in two volumes (Pileckis and Monsevičius 1995, 1997). The introductory section provides a general overview of the main features of the territory of Lithuania, the origins and formation of the beetle fauna and their conservation, the faunistic investigations in Lithuania to date revealing the most important stages of the faunistic research process with reference to the most prominent scientists, an overview of their work, and their contribution to Lithuanian coleopteran faunal research.Species recorded in Lithuania by some authors without reliable evidence and requiring further confirmation with new data are presented in a separate list, consisting of 183 species. For the first time, analysis of errors in works of Lithuanian authors concerning data on coleopteran fauna has been conducted and these errors have been corrected. All available published and Internet sources on beetles found in Lithuania have been considered in the current study. Over 630 literature sources on species composition of beetles, their distribution in Lithuania and neighbouring countries, and taxonomic revisions and changes are reviewed and cited. An alphabetical list of these literature sources is presented. After revision of public beetle collections in Lithuania, the authors propose to remove 43 species from the beetle species list of the country on the grounds, that they have been wrongly identified or published by mistake. For reasons of clarity, 19 previously noted but later excluded species are included in the current checklist with comments. Based on faunal data from neighbouring countries, species expected to occur in Lithuania are matnioned. In total 1390 species are attributed to this category and data on their distribution in neighbouring countries is presented. Completion of this study provides evidence that the Lithuanian coleopteran fauna has yet to be completely investigated and it is estimated that approximately 28 % of beetle species remain undiscovered in Lithuania. More than 85% of beetle species expected for Lithuania have been found in the following families: Cerylonidae, Geotrupidae, Haliplidae, Kateridae, Lycidae, Lucanidae, Mycetophagidae, Scarabaeidae and Silphidae. In families with few species such as Alexiidae, Boridae, Byturidae, Dascilidae, Drilidae, Eucinetidae, Lampyridae, Lymexilidae, Megalopodidae, Nemonychidae, Nosodendridae, Noteridae, Orsodacnidae, Pyrochroidae, Pythidae, Psephenidae, Rhysodidae, Sphaeritidae, Sphaeriusidae, Sphindidae, Stenotrahelidae and Trogidae, all possible species have already been discovered. However in some beetle families such as Aderidae, Bothrideridae, Eucnemidae, Laemoploeidae, Mordellidae, Ptiliidae, Scraptidae and Throscidae less than 50% of all possible species are known. At present the beetle species recorded in Lithuania belong to 92 families, with species from 9 other families such as Agyrtidae, Biphylidae, Deradontidae, Mycteridae, Ochodaeidae, Phleophilidae, Phloeostichidae, Prostomidae, Trachypachidae are expected to be found.A bibliography and a index of subfamily and genus levels are provided. The information published in the monograph will serve to further faunistic and distribution research of beetles and will help to avoid confusion in the identificatation of coleopteran fauna of Lithuania.