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Ungulate Taxonomy by Colin Groves and Peter Grubb (1 Viewer)


Well-known member
United States
Debated which forum to put this on, but given that I feel it's more of interest here than necessarily the books section, and because I am hoping to slowly add the revised species level phylogeny in follow up comments, I figured I should post here.

First off, this is not a popular press book; it's a fairly technical species by species scientific treatise. There are no illustrations at all from what I recall, but plenty of tables and anatomical terminology. For the most part, most of the information is focused on taxonomy, and there is very little behavioral, conservation, etc information, although some stuff is scattered in there.

The genesis of this book was essentially the recent passing of the influential Peter Grubb, who skillfully managed to publish authoritative taxonomic revisions in the scientific literature all the while working as a high school teacher. Colin Groves essentially uses this book to review his and Grubb's papers (They were often coauthors, besides friends), as well as a polishing up and publishing many uncomplete manuscripts Grubb had worked on before his passing.

The book covers all Artiodactyl and Perissodactyl families; Elephants, hyraxes, and aardvarks are not discussed. The first 10 pages provide a brief introduction, including describing the species concept they use, the history of ungulate taxonomy, and the impact the new taxonomy has on biogeography and conservation. This is a well written introductory bit that I think most layman would appreciate.

The rest of the book is a family by family break down and review of subfamily, genus, species, and subspecies taxonomy, including their revisions. If you have seen the Bovidae chapter of Handbook of Mammals of the World, you know what to expect, with the biggest impact probably effecting pigs, deer, giraffes, and of course the bovids.

The writing and quality of the different sections is fairly uneven. Some heading give Common names and detailed range information; some don't. Usually the authors cite other papers, but some entries advise readers to check out Wikipedia links (no, really...). While I understand the lack of illustrations, range maps, even if they only only contained locality dots, would be incredibly helpful. Thankfully the Bovidae chapter, the worst victim of lack of range information, can be crosschecked with HMW. Often keywords are thrown out at the beginning of a section, and then completely ignored. At times one gets a rather "rush job" feeling for the writing, which overall could use further polishing

The same could go for the reasoning behind many splits. The authors rely on a fairly small number of morphological measurements to support their taxonomic revisions, and often many species are represented by small sample sizes. No doubt many of these splits are valid and will receive further support, but one does wonder if a larger sample size or more exhaustive look might not produce different results. I will be interested in seeing how well many of these changes are accepted.

I should also mention that this is by far not a final revision of artiodactyl taxonomy. The authors back away from some groups, and little revision takes place. New World Deer and to a lesser extent pigs are probably the greatest victims of this, though the disregard for Elephants (considered ungulates by almost everyone) is worse. It's actually no terribly obvious how many species the authors recognize in the former two groups.

Anyway, this is my take on things. I will be adding the species level taxonomy in subsequent posts on this thread, for anyone interested but without access to this book.

Please note that for brevity sake I am not going to list their subspecies classification. Also I will only list ranges when they are especially confusing

Tarpan (Equus ferus) - extinct
Przewalski's Horse/Takhi (Equus przewalskii)
African Ass (Equus africanus)
Kiang (Equus kiang)
Onager/Asian Wild Ass (Equus hemionus)
Khur/Indian Wild Ass (Equus khur)
Achdari/Syrian Wild Ass (Equus hemippus) - extinct
Cape Mountain Zebra (Equus zebra)
Hartmann's Zebra (Equus hartmannae)
Grevy's Zebra (Equus grevyi)
Plains Zebra (Equus quagga)

South American Tapir (Tapir terrestris)
Mountain/Woolly Tapir (Tapir pinchaque)
Baird's Tapir (Tapirella bairdii)
Malay Tapir (Acrocodia indica)

Indian/Greater One-horned Rhino (Rhinoceros unicornis)
Javan/Lesser One-horned Rhino (Rhinoceros sondaicus)
Sumatran Rhinoceros (Dicerorhinus sumatrensis)
Black Rhinoceros (Diceros bicornis)
Southern White Rhinoceros (Ceratotherium simum)
Northenr White Rhinoceros (Ceratotherium cottoni)
Thanks Morgan,
is the Equus quagga in this treatment also an extinct species, or which extant subspecies is included?

Artiodactylidae: Camels - Hippos

Also worth noting is that domestic species are not covered (though a complete list is given in the Introduction material)

Guanaco (Lama guanicoe)
(Lama vicugna) - no common name given (?Chilean Vicuna) -W Bolivia, NW Argentina through Chile
(Lama mensalis) - no common name given (?Peruvian Vicuna) -SE Peru, W Bolivia, and NE Chile

I culled the range information from other sources...this book does not clearly state ranges

Wild Bactrian Camel (Camelus Ferus)

Buru Babirusa (Babyrousa babyrussa)
Togean Babirusa (Babyrousa togeanensis)
North Sulawesi Babirusa (Babyrousa celebensis)
(?Eastern Giant Forest Hog) Hylochoerus meinertzhageni - east of Albertine Rift)
(?Central? Giant Forest Hog) Hylochoerus rimator - Central Africa
(?Western Giant Forest Hog) Hylochoerus ivoriensis - West Africa

Again no common names given for the above animals

Pygmy Hog (Porcula salvania)
Desert Warthog (Phacochoerus aethiopicus)
Common Warthog (Phacochoerus africanus)
Bushpig (Potamochoerus larvatus)
Red River Hog (Potamochoerus porcus)

The authors consider the below group to be a work in progress essentially

?Common/Western Wild Boar (Sus scrofa) - Europe, North Africa, and Middle East
?Corsican Wild Boar (?Sus meridionalis)- Corsica and Sardinia, maybe Southern Spain - authors undecided on validity of taxon
?(Sus nigripes) - known from one Chinese specimen, unexamined by authors
Siberian Wild Boar (?Sus sibiricus) - mongolia and Transbaikalia - unexamined by authors who are unclear on validity of taxon
Japanese Wild Boar - (Sus leucomystax) - Japan
Formosan Wild Boar - (Sus taevanus) - Taiwan
Ryuku Wild Boar - (?Sus riukiuanus) - Ryuku Islands - needs more comparison with Formosan Wild Boar
Central Asian Wild Boar (Sus davidi) - Iran, east of Zagros mountains - Pakistan
Indian Wild Boar (Sus cristatus) - India
?Heude's Wild Boar (Sus moupinensis) - Burma-China
Southern Chinese Wild Boar (Sus chirodontus) -South Central China
Manchurian Wild Boar (Sus ussuricus) - Far East
Banded Pig (Sus vittatus) - SE Asia

Yeah...that's a 9-12 way split of Wild Boar...

Sulawesi Warty Pig (Sus celebensis)
Javan Warty Pig (Sus verrucosus)
Bawean Warty Pig (Sus blouchi)
Bearded Pig (Sus barbatus)
Palawan Wild Pig (Sus ahoenobarbus) - Palawan, Balabac, and the Calamianes
Philippine Wild Pig (Sus philippensis) - Luzon, Mindanao, and offshore islands
Mindoro Wild Pig (Sus oliveri) - Mindoro
Visayan Warty Pig (Sus cebifrons)

White-lipped Peccary (Tayassu pecari)
Southern Collared Peccary (Pecari tajacu) - Mato Grosso, N Argentina, and Paraguay
Northern Collared Peccary (Pecari angulatus) - Northern Mexico and USA
?Central American Peccary (Pecari crassus) - rainforest zone from Southern Mexico to Ecuador and Columbia, presumably further south, although the text suggests a 4th unnamed species here? (confused)
Chaco Peccary (Catagonus wagneri)

Common Hippopotamus (Hippopotamus amphibius)
Western Pygmy Hippopotamus (Choeropsis liberiensis)
Niger Delta Pygmy Hippopotamus (Choeropsis heslopi) - probably extinct
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The Quagga is considered an extinct subspecies of the Plains Zebra, and the odd patterning probably the end result of clinal variation. This is actually a pretty non controversial stance
Great stuff Morgan...

Currently trembling though cos I might have to repaint some plates.... and my HMW still hasn't arrived!!
Mouse-Deer, Musk Deer, Pronghorn, and Giraffes


Note no common names listed, so going off of Wikipedia (which in turns goes off of MSW3). At any rate, no novel splits recognized, although the authors do conclude more work needs to be done, and there are several oddball specimens they don't know where to place

Greater Mouse-Deer (Tragulus napu)
Philippine Mouse-Deer (Tragulus nigricans)
Vietnam Mouse-Deer (Tragulus versicolor)
Java Mouse-Deer (Tragulus javanicus)
Lesser Mouse-Deer (Tragulus kanchil)
Williamson's Mouse-Deer (Tragulus williamsoni)
Indian Spotted Chevrotain (Moschiola indica)
Sri Lankan Spotted Chevrotain (Moschiola memina)
Yellow-striped Chevrotain (Moschiola kathygre)
Water Chevrotain (Hyemosuchus aquaticus)


Same as the above

Siberian Musk Deer (Moschus moschiferus)
Alpine Musk Deer (Moschus chrysogaster)
White-bellied Musk Deer (Moschus leucogaster)
Kashmir Musk Deer (Moschus cupreus)
Black Musk Deer (Moschus fuscus)
Anhui Musk Deer (Moschus anhuiensis)
Dwarf Musk Deer (Moschus berezovskii)


Pronghorn (Antilocapra americana)


Nubian/Rothschild's Giraffe (Giraffa camelopardalis)
Reticulated Giraffe (Giraffa reticulata)
Kordofan Giraffe (Giraffa antiquorum)
West African Giraffe (Giraffa peralta)
Masai/Vine-leaf Giraffe (Giraffa tippelskirchi)
Luangwa Giraffe (Giraffa thornicrofti)
Cape Giraffe (Giraffa giraffa)
Angolan Giraffe (Giraffa angolensis)
Okapi (Okapia johnstoni)
Deer (New World)


White-tailed Deer (Odocoileus virginianus)
Black-tailed Deer (Odocoileus columbianus)
Mule Deer (Odocoileus hemionus)

Note that the authors do not fully revise this genus; they call for further work and suggest future splits of South American and Central American forms, at the very least. The generic classification also needs to be fully revised

Marsh Deer (Blastocerus dichotomus)
Pampas Deer (Ozotocerus bezoarcticus)
Taruka (Hippocamelus antisensis) - Andean Peru to Northern most Chile
Huemel (Hippocamelus bisulcus) - Andean south Chile and Argentina
Small Red Brocket (Mazama bororo) - Atlantic forest of Brazil
Pygmy Brocket (Mazama nana) - SE Brazil, NE Argentina - easternmost Paraguay
? Red Brocket (Mazama whitelyi) - South Peru
Common Red Brocket (Mazama americana) - Guyanas, S and E Venezuala, NE Brazil
? Red Brocket (Mazama jucunda) - Sao Paulo to Rio Grande do sul
? Red Brocket (Mazama zamora) - SE Colombia, Ecuador east of Andes, and NE Peru
? Red Brocket (Mazama zetta) - Inter-Andean valleys in Columbia
Gualea Red Brocket (Mazama gualea) - Ecuador west of the Andes, possibly SW Colombia
Central American Red Brocket (Mazama temama) - SE Mexico to NW Colombia
Trinidad Red Brocket (Mazama trinitatis) - Trinidad
Merida Brocket (Mazama bricenii) - Andes of W Venezuala and neighboring Colombia
Little Red Brocket (Mazama rufina) - Premontane forest of S Colombia and Ecuador
Venezualan Brown Brocket (Mazama cita) - Northern Venezuala
Gray Brocket (Mazama gouazoubira) - Southern Brazil to Northern Argentina and Uruguay
? Brown Brocket (Mazama murelia) - SE Colombia and Ecuador east of Andes
Amazonian Brown Brocket (Mazama nemorivaga) - Guyanas, N Brazil, SE Venezuala
? Brown Brocket (Mazama permira) - Isla San Jose, Panama
? Brown Brocket (Mazama sanctaemartae) - Northern Colombia
? Brown Brocket (Mazama superciliaris) - Central and Eastern Brazil, from Amazonas to Serra dos Parecis in Mato Grosso; Espiritu Santo
? Brown Brocket (Mazama tschudii) - Peruvian Cordillera
Rufous Brocket (Mazama rondoni) - Atlantic Brazil
Yucatan Brown Brocket (Mazama pandora) - Yucatan
Dwarf Brocket (Mazama chunyi) - premontane forests of Andean Peru/Bolivia

Several new taxa were also suggested, but lacked available names. I couldn't find any common names described for most of these poorly known deer

Southern Pudu (Pudu puda)
Northern Pudu (Pudu mephistopheles)
Caribou/Reindeer (Rangifer tarandus)
Cervidae (Old World)

Will do the Bovidae tomorrow!

Western Roedeer (Capreolus capreolus)
Siberian Roedeer (Capreolus pygargus)
Chinese Water Deer (Hydropotes inermis)
Elk (Alces alces) - Old World
Moose (Alces americana) - New World + eastern Siberia
Tufted Deer (Elaphodus cephalophus)
Southern Red Muntjac (Muntiacus muntjak) - Java, Bali, Lombok, Borneo, Bangka, Lampung, coastal Sumatra, and Malay peninsula
Northern Red Muntjac (Muntiacus vaginalis) -Nepal, east India to W Burma
?Malabar Muntjac (Muntiacus malabaricus) - Sri Lanka and W Ghats
? Muntjac (Muntiacus aureus) - NW/C India, lower Chindin in Burma
? Muntjac (Muntiacus nigripes) - Hainan, Northernmost Vietnam
Hairy-fronted Muntjac (Muntiacus crinifrons) - W Zheijang, SE Anhui, to Fujian
Gongshan Muntjac (Muntiacus gongshanensis) - Gaoligong and Bilou Mtns, China
Fea's Muntjac (Muntiacus feae) - S Thailand
Giant Muntjac (Muntiacus vuquangensis) - Vietnam, Central Laos
Roosevelt's Muntjac (Muntiacus rooseveltorum)
Leaf Muntjac (Muntiacus putaoensis) N Burma)
Truong Son Muntjac (Muntiacus truonsonensis) - Truong son Mtns, Vietnam
Pu Hoat Muntjac (Muntiacus puhoatensis) - Pu Hoat region, Vietnam
Chinese Muntjac (Muntiacus reevesi) - China and Taiwan
Bornean Yellow Muntjac (Muntiacus atherodes) - Borneo
Sumatran Muntjac (Muntiacus montanus) - Sumatran highlands

Common Fallow-deer (Dama dama)
Mesopotamian Fallow-deer (Dama mesopotamica)
Indian Hog-deer (Axis porcinus) - Sri Lanka, Sind E to Nepal and NE India
Indochina Hog-deer (Axis annamiticus) - Indochina?
Calamanian Deer (Axis calamianensis)
Bawean Deer (Axis kuhli) -
Western Swamp Deer/Barasingha (Rucervus duvaucelii) - N India and Nepal
Eastern Swamp Deer (Rucervus ranjitsinhi) - Assam
Hard-ground Barasingha (Rucervus branderi) - Central India
Schomburgk's Deer (Rucervus schomburgki) - extinct
Sangai/Manipur Eld's Deer (Panolia eldii) - India
Thamin (Panolia thamin) - Burma and W Thailand
Eastern Eld's Deer (Panolia siamensis) - E Thailand, Cambodia, S Laos, Hainan, ?Vietnam
Pere David's Deer (Elaphurus davidianus)

Now time for the Red deer...

West European Red Deer (Cervus elaphus) - Western Europe
East European Red Deer (Cervus pannoniensis) - Austria, Bulgaria, Hungary, Yugoslavia, Romania, and Turkish Thrace
Turkish/Persian Red Deer (Cervus maral) - Middle East
Barbary Stag (Cervus corsicanus) - North Africa, Corsica and Sardinia
Yarkand Stag (Cervus yarkandensis) - Tarim Basin, Xinjiang
Bactrian Stag (Cervus bactrianus) - Syr Darya and Amu Darya Basins
Kashmir Stag (Cervus hanglu) - Kashmir
Tibetan Shou (Cervus wallichii) - S Tibet into Sikkim
Sichuan Shou (Cervus macneilli) - Sichuan and Gansu, China
Alashan Stag (Cervus alashanicus) - Mongolia?
Manchurian Wapiti (Cervus xanthopygus) - Yakutia, Amuria, and Manchuria
American Wapitit (Cervus canadensis) - North America, Tianshan and Altai

White-lipped Deer (Cervus albirostris)

Sika gets carved up too

Japanese Sika (Cervus nippon) - Japan and Ryuku islands south of Biwa ko
Hokkaido Sika (Cervus aplodontus) - Japan north of Biwa ko (Hokkaido and Northern Honshu)
Tsushima Sika (Cervus pulchellus) - Tsushima island
Manchurian Sika (Cervus hortulorum) - N China and Russian Far East
Sichuan Sika (Cervus sichuanicus) - Sichuan
Formosan Sika (Cervus taiouanus) - Taiwan
Vietnamese Sika (Cervus pseudaxis) - Vietnam and South China

Prince Alfred's Deer (Cervus alfredi) - Lyte, Cebu, Guimaras, Negros, and neighboring Philippine islands
Indian Sambar (Cervus unicolor) - Sri Lanka and mainland India
Southeast Asian Sambar (Cervus equinus) - Bengal and Assam, NE into S China and Taiwan, and SE to Bornea and Malay Penisula
Philippine Deer (Cervus mariannus) - Luzon, Mindanao, Basilan, and introduced to Guam
Mindoro Deer (Cervus barandanus) - Mindoro
Mindanao Mountain Deer (Cervus nigellus) - highlands of Mindanao
Rusa (Cervus timorensis) - Java and Bali, introduced as far east as Timor and Moluccas
Wow! However, although it probably reflects reality, I'm not sure how useful a list with so many questioned species is. Will have to cast my eyes on a copy at some stage.
This is return of 19. century approach. Then people recognized lots of similar species (over 20,000 species of mammals).

Ungulates, like most animals, change geographically in larger clines plus lots of local varieties. If a hunter or scientists picks distant dots in the species range, will see populations with many small but distinct differences. It is conceivable that he can give them different names. But his follower will discover that he cannot see these species. Between dots there are intermediates or new distinct populations. Somebody having good overview of a species along whole geographic range will note that these differences are clinal and only the broad name is useful.

This happened in 19. century, when hunters shot an antelope here, another there, and many local species were named. As the knowledge of the world's fauna became more complete, it became obvious that there are intermediates, and only broad name is useful.

Perhaps significantly, often the less data exist, the more Groves & Grubb split. So few splits in Europe or North America, because they are so well known that everybody realizes that local varieties mix. But brocket deer from South American rainforest or gazelles from politically inaccessible Middle East are split to the will.

It is interesting to see artifact of politics and economic development. If a region is politically or logistically inaccessible to scientists, it turns up as a barrier artificially creating new species on the sides. Also, many continious ungulate populations were hunted out in recent decades. This left small and inbred groups in national parks and reserves. People see each of this populations as distinct, because intermediates are gone. So there is a situation that the less biodiversity, the more is species.

The lesson from early 20. century is that splitting was abandoned. It was not because of arguments of theoretical BSC/PSC debate (these concepts were yet unknown) but because it was impossible in practice.

It cannot be applied to other groups, even vertebrates (10,000s of frogs, lizards or rodents, anyone?), creates many hybrid populations, many populations where species cannot be determined, and too small differences in general.
A load of armchair ticks for me in there ( if I were to accept the recommendations! ). Some rather excessive splits though, IMO, and Roedeer, Fallow-deer, Hog-deer, Mouse-deer ???????? ( but not Red-deer, Mule-deer or, more importantly, Musk-deer! ) Are we going to end up with more stupid distortions of English because of a lack of knowledge of grammar and etymology? Roe Deer and Fallow Deer are names of deer that go back well before formal taxonomy and to infer that they are, somehow, not 'real' deer by combining / hyphenating the names is.................... :C I know its all been hashed over with bird names but...........:-C

I think the reason there are so few new world splits isn't because the fauna is so well known (it's not), it's because the authors have a bias towards old world taxa, especially given that Peter Grubb was based out of England, while I think Colin Groves has had multiple positions but none in the US. Also at least some splits are apparently of taxa in sympatry or with sharp breaks in any sort of morphocline. While I think some of the splits are excessive, I actually think mammal taxonomy is over conservative, and a break up of some of these groups is warranted. We've butted heads in the past on this however.

Interestingly enough, the Mazama splits I actually found to be pretty persuasive. If you look at the distribution, most are found in areas of with a high number of endemic birds, so it makes sense that different Andean Slopes, Santa Marta Mtns, Atlantic Rainforest, etc would probably have endemic forms of deer, especially given the the fact that very few people have really worked on this group and new species are still being found.

Chris, the common names are not a purpose of this book, and the author makes no real effort to standardize them (half the species have no common name listed). Feel free to pick or choose what you want to call them, all the authors care about is making sure you use the right species/genus name :p
One other point: as I listed, questionable taxa were placed with a ? next to the scientific name. Well supported taxa in their opinion, but species which I could find no common name, have a ? by the "common name" That I think mostly occurs with pigs.
The Quagga is considered an extinct subspecies of the Plains Zebra, and the odd patterning probably the end result of clinal variation. This is actually a pretty non controversial stance

Thanks, but I am probably a little naive about this one. Mammals of the world 3 treats E. quagga as an extinct monotypic species, I believe. E. burchellii has six subspecies in this treatment. I think the older treatment was to lump these to one species with seven subspecies. So that is why I am asking which other subspecies are included in Plains Zebra in the new book? (If I knew more about mammals, this would probably be obvious to me, but isn't).

No problem...didn't realize that MSW3 considered Quagga a monotypic species. I can give you a run down of subspecies if you want tomorrow, but after an entire evening spent looking at phylogenies and teeth, I mostly want to go to bed now!
Just because you were curious, here are the subspecies listed for Plains Zebra

Quagga (E.q. quagga)
Burchell's Zebra (E.q. burchellii)
Chapman's Zebra (E. q. chapmani)
Crawshay's Zebra (E. q. crawshayi)
Grant's Zebra (E. q. boehmi)
Half-maned Zebra (E. q. borensis)
Indian Hog-deer (Axis porcinus) - Sri Lanka, Sind E to Nepal and NE India
Indochina Hog-deer (Axis annamiticus) - Indochina?
Calamanian Deer (Axis calamianensis)
Bawean Deer (Axis kuhli) -

What happened to Chital (Axis axis)?

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