Fauna och Flora, Vol. 56, Nº 4, 2011, pp.42-45

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All about Hoofed Mammals in HMW, Vol. 2

Ph. Dr. Torbjörn Ebenhard, Swedish BiodiversityCentre, Swedish University of Agricultural Sciences

Wilson, D. E. & Mittermeier, R. A. (eds.) 2011. Handbook of the Mammals of the World. Vol. 2. Hoofed mammals. Lynx Edicions, Barcelona.

The second volume of the mammoth encyclopaedia Handbook of the Mammals of the World has recently been published. It covers the ungulates and a number of other mammal groups which were until recently regarded as their close relatives, but have now been transferred to other parts of the mammal genealogical tree. The subdivision of the mammal groups between and within the eight planned volumes thus seems tube governed by commercial concerns rather than recent systematics. In this volume the odd-toed and even-toed ungulates are presented together with the Aardvark, the hyraxes and the elephants, despite the fact that the three latter are currently placed within the clad Afrotheria along with sea cows (sirenians),tenrecs and golden moles, which will be presented in a forthcoming volume. The pangolins are also included in this volume, although their closest relatives are the carnivores. The whales are the direct descendents of even-toed ungulates, but they will none the less be presented in Volume 4, together with all other aquatic mammals.

Volume 2 covers 17 mammal families with atonal of 413 species. The only Swedish species presented in this volume are the Eurasian Wild Pig, Common Fallow Deer, Western Red Deer, Western RoeDeer, Moose (Elk), Reindeer and Muskox – but not the Mouflon, which is regarded as a feral sheep. Domesticated mammals are usually not presented, or at least not given full presentations. They are usually mentioned in the family presentations, along with a brief survey of their role in relation to man. In some cases (e.g. the domestic pig, the Mithan and the Bali Cattle) they are regarded as mere domesticated varieties of their respective wild original species, but more usually they are treated as separate species. One notable exception from the principle of not treating domestic species is made in the case of the camelids,where also the Dromedary Camel, the Llama and the Alpaca are given full species presentations. Each family is given a general presentation covering systematics, morphology, habitat selection, ethology and communication, food and feeding, reproduction, movements and home range, social structure, relationship with humans, status and conservation, followed by a selected bibliography. These family presentations range from six pages (Aardvark) to 128pages (bovids). They are followed by presentations of each included species. The headings are roughly the same in the species presentations, with additional information about subspecies, distribution and diagnostic characters. The text is written by a team of 29 international experts of zoology, ecology, ethology, veterinary medicine, conservation biology and game management, headed by Don Wilson and Russell Mittermeier (American taxonomist and primatologist/conservation biologist, respectively). Despite the massive format (32×25 cm, 885 pages, 4.4 kg), this is a work of popular science. The language is relatively advanced, but not strictly scientific. Unfortunately, there is no glossary explaining technical terms. Especially the family presentations are, however, easy to read, whereas the species presentations are somewhat more compact and crammed with facts. References are not given for each individual piece of information, but there is bibliography at the end of every chapter/species presentation.

Almost every opening of the family presentations is illustrated by three or four large colour photos, in total 663 top quality photos. The photos are more than straightforward species portraits; they illustrate the characteristic behaviour, reproduction and habitats of the species. Almost all of them depict wild specimensat well-specified sites, and all are identified to species. All genera except one (Ammodorcas), and179 out of the 413 species are presented by at least one photo. It may seem somewhat mediocre to only show photos of a bit less than half of the presented species, but the photo material of this book is none the less without parallel in modern book production– both in terms of quality and quantity. In addition, each species is depicted in one of the 56 exquisite colour plates. On average, 8 species are presented per plate (both sexes, often also subspecies). The plates are produced by Toni Llobet; an autodidact, Spanish illustrator, who has been contracted for all eight volumes. He utilises digital tools enabling him to work swiftly – an absolute necessity if he is to cover over6 000 species. However, the most daunting aspect of his formidable task is maybe not depicting all the species, but learning the characteristic features of each and every one of them. This is not primarily an identification handbook, even if it serves also that purpose with regard to the larger species. When it comes to the smaller mammals it will, however, be difficult to demonstrate the difference between the species by the type of illustrations used in the first two volumes. To do that, detailed illustrations of e.g. skulls and teeth would be required. So far (in the first two volumes), facts about inner anatomy; skulls, skeletal constructions, teeth, internal organs etc. are given brief verbal descriptions in the family presentations, but no illustrations. Also physiology, diseases and parasites are only treated summarily. In the introductory chapter of Volume 1, the editors stated that the Handbook would follow the conventional mammal systematics of Wilson & Reeder(2005), which was perfectly true of the first volume. In Volume 2, however, this principle already seems to have been abandoned, especially with regard to the bovine family (Bovidae). The main author of this chapter, Colin Groves, instead follows the new systematic of this family, which was scientifically presented by himself and Peters Grubbs as late as December this year. Groves is known to advocate a phylogenetic species concept, where a species is defined as the smallest population or group of populations displaying diagnostic, genetically based differences compared to other populations. To put it simply; a species is a population deviating from other populations with regard to at least one character. This species concept differs markedly from the so called biological species concept prevalent during the latter half of the 20thcentury, according to which two separate species must be reproductively isolated from each other in order to be regarded as such (which is not a pre-requisite according to the phylogenetic species concept). The effect of switching from the old to the new species concept is a dramatic increase in the number ofspecies,as many of the former species become subdivided into several units. The number of bovines has thus increased from 143 in Wilson & Reeder (2005)to 279 in this volume of the Handbook. The editors have, however, not made sure that all authors apply the same species concept. The Giraffe is, for instance, still treated as a single species with a number of distinct subspecies, with the motivation that there is no reproductive barrier between the subspecies.

Many well-known species, e.g. the Hartebeest, the Bushbuck, the Sitatunga and the African Buffalo, have been split. In the case of the Klipspringer, the number of currently recognised species actually exceedsthe number of former subspecies. The sheepgenus (Ovis) has acquired 16 new species, and 13new species have been added to the genus Gazella. Several new genera have also been established, encompassing the species which were formerly regarded as subspecies within a single species. The new genera Ammelaphusand Strepsiceros, for instance, encompass two and four species, respectively, of what was formerly the two Kudu species in the genus Tragelaphus. Such taxonomical changes are not, strictly speaking, based on new knowledge, but rather on a revaluation of what a species is. The advocates of the new species concept maintain that it gives a much clearer picture of the extant biodiversity, whereas the detractors regard it more as a kind of taxonomical mischief. A common argument in the conservation debate is that it’s important to give all distinctive population the status of separate species because this makes it easier to motivate their protection and preservation. As pecies is regarded as an indispensible entity, whereas a subspecies tends to be viewed as a local variety and neglected both by the red-listing organisations and the authorities who should protect it. This is not valid reason for making taxonomical revisions and, besides, there is no natural (or juridical) law stating that subspecies have to be given a lower status in the contexts of red-listing or conservation legislation.

The new Handbook volume does, however, also present taxonomical novelties that are actually based on new biological knowledge. Two new genera, Arabitragus and Nilgiritragus, have been defined in accordance with the insight that each of the three tahr species formerly constituting the genus Hemitragusare, in fact, more closely related to other sheep and goat species than to each other. A number of recently described species – e.g. the Saola and three muntjac species (Muntiacus puhoatensis, M. truongsonensis and M.putaoensis) – are, of course, also presented. According to Lynx Edicions, the purpose of the Handbook is to describe all extant wild mammal species and their conservation status, and to promote the preservation of them and their habitats. Does Volume2 of the Handbook, then, live up to these ambitions? Is it a reliable reference work? Indeed, it is! It is an unprecedented overview of the mammals of the world and an impressive top quality fact collection. Having said that, there are, of course, always minor errors. I have, for instance, spotted three miss-spellings of scientific names (one of them being the very first one mentioned in the introductory chapter)!

Can the Handbook contribute to the preservation of the mammal species and their habitats? That is less certain. To all those who get the chance to use it in their education or their work, it provides excellent ammunition in the struggle for mammal conservation. But considering the voluminosity and the high price of the entire Handbook, only those who are already the most dedicated conservationists and mammal enthusiasts are likely to buy the entire Handbook. My hope is that it will be made accessible to a wider audience through schools and public libraries. The range and population size of most large mammal species is decreasing, and many of them are directly threatened by extinction. In the presentation of the Javan Rhinoceros, it is mentioned that what may have been the last specimen in Vietnam was found dead in 2010. Very recently it was confirmed that this was the case; the Javan Rhinocerosis extinct on the Asian mainland, and currently only some 40 individuals survive on Java. The distribution maps for the two African rhinoceros species are very vague, covering most of the continent, in order not to reveal the exact location of the last individuals. It does, however, seem more than a bit far-fetched that African poachers would use this Handbook volume to localise their prey. It would probably have been wiser to demonstrate how very limited their current distribution is, thereby presenting a more correct picture to the readers.

There are six more volumes to publish before the Handbook is complete. A bit more than two years passed between the first two volumes, but no indication is given of the future publication rate. Another question is how the editors will manage to cover all the remaining species. This volume comprises a bit more than 400 species, and in Handbook of the Birds of the World the maximum number of species per volume was a bit over 800 (del Hoyo, Elliot & Christie2003). Volume 6 is intended to cover all the over2 000 rodent species! Irrespective of how this problem is solved, and of the publication rate, I will really look forward to each new publication date! Even if I would not acquire any other mammal books for the rest of my professional life, I would still buy all the remaining volumes of Handbook of the Mammals of the World.

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