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Ungulate taxonomy revisited: the evidence for the splits of G&G

Discussion in 'Wildlife & Nature Conservation' started by lintworm, 1 Jul 2017.

  1. TeaLovingDave

    TeaLovingDave Moderator Staff Member 10+ year member

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    And, more notably, prehistoric hybridization between burmanicus and Kouprey - this being the reason for the unclear taxonomic status of the latter taxon for a time.
     
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  2. lintworm

    lintworm Well-Known Member 15+ year member

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    The evidence for the splits of Groves & Grubbs


    The Bovid family tree traditionally looked as follows:

    BOVIDAE

    Bovinae

    Boselaphini

    Boselaphus
    Nilgai (Bosephalus tragocamelus)

    Tetracerus
    Four-horned antelope (Tetracerus quadricornis)

    Bovini

    Syncerus
    African buffalo (Syncerus caffer)

    Bubalis
    Asian wild buffalo (Bubalis arnee)
    Tamaraw (Bubalis mindorensis)
    Lowland anoa (Bubalis depressicornis)
    Mountain anoa (Bubalis quarlesi)


    Bos
    Gaur (Bos gaurus)
    Banteng (Bos javanicus)
    Kouprey (Bos sauveli)
    American bison (Bos bison)

    Wisent (Bos bonasus)
    Yak (Bos mutus)

    Pseudoryx
    Saola (Pseudoryx nghetinhensis)

    Tragelaphini

    Tragelaphus
    Lesser kudu (Tragelaphus imberbus)
    Bushbuck (Tragelaphus scriptus)
    Nyala(Tragelaphus angasii)
    Eland (Tragelaphus oryx)
    Giant eland (Tragelaphus derbianus)

    Greater Kudu (Tragelaphus strepsciceros)
    Mountain nyala (Tragelaphus buxtoni)
    Sitatunga (Tragelaphus spekii)
    Bongo (Tragelaphus euryceros)


    Antilopinae

    Neotragini

    Neotragus
    Royal antelope (Neotragus pygmaeus)
    Bates's pygmy antelope (Neotragus batesi)


    Nesotragus
    Suni (Nesotragus moschatus)


    Procaprini

    Procapra
    Tibetan gazelle (Procapra picticaudata)
    Przewalski's gazelle (Procapra przewalskii)
    Mongolian gazelle (Procapra gutturosa)



    Raphicerini

    Raphicerus
    Sharpe's grysbok (Raphicerus sharpei)
    Cape grysbok (Raphicerus melanotis)
    Steenbok (Raphicerus campestris)



    Madoquini

    Madoqua
    Salt's dikdik (Madoqua saltiana)
    Silver dikdik (Madoqua piacentinii)

    Guenther's dikdik (Madoqua guentheri)
    Kirk's dikdik (Madoqua kirkii)


    Ourebini

    Ourebi
    Oribi (Ourebia oribi)


    Antilopini

    Saiga
    Saiga (Saiga tartarica)

    Litocranius
    Gerenuk (Litocranius walleri)

    Ammodorcas
    Dibatag (Ammorodorcas clarkei)

    Antilope
    Blackbuck (Antilope cervicapra)

    Antidorcas
    Sprinbok (Antidorcas marsupialis)

    Eudorcas
    Thomson's gazelle (Eudorcas thomsoni)
    Mongalla gazelle (Eudorcas albonotata)
    Heuglin's gazelle (Eudorcas tilonura)
    Red-fronted gazelle (Eudorcas rufifrons)


    Nanger
    Grant's gazelle (Nanger granti)
    Dama gazelle (Nanger dama)
    Soemmerring's gazelle (Nanger soemmerringi)


    Gazella
    Dorcas gazelle (Gazella dorcas)
    Saudi gazelle (Gazella saudiya)
    Slender-horned gazelle (Gazella leptoceros)
    Cuvier's gazelle (Gazella cuvieri)

    Sand gazelle (Gazella marica)
    Speke's gazelle (Gazella spekei)
    Goitered gazelle (Gazella subgutturosa)
    Arabian gazelle (Gazella arabica)
    Indian gazelle (Gazella bennetti)


    Oreotragini

    Oreotragus
    Klipspringer (Oreotragus oreotragus)


    Reduncini

    Redunca
    Common Reedbuck (Redunca arundinum)
    Bohor reedbuck (Redunca bohor)
    Mountain reedbuck (Redunca fulvorufula)

    Kobus
    Kob (Kobus kob)
    Puku (Kobus vardoni)
    Southern lechwe (Kobus leche)
    Nile lechwe (Kobus megaceros)
    Waterbuck (Kobus ellipsiprymnus)

    Pelea
    Grey rhebok (Pelea capreolus)


    Aepycerotini

    Aepyceros
    Impala (Aepyceros melampus)


    Cephalophini

    Philantomba
    Blue duiker (Philantomba monticola)
    Maxwell's duiker (Philantomba maxwelli)
    Walter's duiker (Philantomba walteri)


    Sylvicapra
    Common duiker (Sylvicapra grimmia)

    Cephalophus
    Bay duiker (Cephalophus dorsalis)
    Yellow-backed duiker (Cephalophus silvicultor)
    Ogilby's duiker (Cephalophus ogilbyi)
    Weyn's duiker (Cephalophus weynsi)
    White-bellied duiker (Cephalophus leucogaster)
    Natal red duiker (Cephalophus natalensis)
    Harvey's duiker (Cephalophus harveyi)

    Black-fronted duiker (Cephalophus nigrifrons)
    Zebra duiker (Cephalophus zebra)
    Jentink's duiker (Cephalophus jentinki)
    Abbot's duiker (Cephalophus spadix)
    Black duiker (Cephalophus niger)
    Ader's duiker (Cephalophus adersi)
    Red-flanked duiker (Cephalophus rufilatus)
    Rwenzori red duiker (Cephalophus rubidus)
    Peter's duiker (Cephalophus callypigus)



    Alcelaphini

    Damaliscus
    Bontebok/Bleskbok (Damaliscus pygargus)
    Topi/Tsessebe/Tiang/Korrigum (Damaliscus lunatus)

    Alcelaphus
    Hartebeest (Alcelaphus busephalus)

    Connochaetes
    Common wildebeest (Connochaetes taurinus)
    White-tailed wildebeest (Connochaetes gnu)

    Beotragus
    Hirola (Beotragus hunteri)


    Hippotragini

    Hippotragus

    Sable antelope (Hippotragus niger)
    Roan antelope (Hippotragus equinus)
    Bluebuck (Hippotragus leucophaeus)


    Oryx
    Beisa oryx (Oryx beisa)
    Gemsbok (Oryx gazella)
    Scimitar-horned oryx (Oryx dammah)
    Arabian oryx (Oryx leucorix)


    Addax
    Addax (Addax nasomaculatus)


    Caprini

    Pantholops
    Chiru (Pantholops hodgsoni)

    Ovibos
    Muskox (Ovibos moschatus)

    Naemorhedus
    Himalayan goral (Naemorhedus goral)
    Chinese goral (Naemorhedus griseus)
    Long-tailed goral (Naemorhedus caudatus)
    Red goral (Naemorhedus baileyi)


    Capricornis
    Sumatran serow (Capricornis sumatraensis)
    Chinese serow (Capricornis milneedwardsi)
    Himalayan serow (Capricornis thar)
    Burmese red serow (Capricornis rubidus)
    Formosan serow (Capricornis swinhoei)
    Japanese serow (Capricornis crispus)


    Ovis

    Argali (Ovis ammon)
    Urial/Mouflon (Ovis vignei)

    Snow sheep (Ovis nivicola)
    Bighorn sheep (Ovis canadensis)
    Dall's sheep (Ovis dalli)


    Ammotragus

    Barbary sheep (Ammotragus lervia)

    Arabitragus
    Arabian tahr (Arabitragus jayakari)

    Nilgiritragus

    Nilgiri tahr (Nilgiritragus hylocrius)

    Rupicapra

    Alpine chamois (Rupicapra rupicapra)
    Pyrenean chamois (Rupicapra pyrenaica)


    Budorcas
    Takin (Budorcas taxicolor)

    Oreamnos
    Rocky Mountain goat (Oreamnos americanus)

    Pseudois
    Blue sheep (Pseudois nayaur)

    Hemitragus
    Himalayan tahr (Hemitragus jemlahicus)

    Capra
    Wild goat (Capra aegagrus)
    Markhor (Capra falconeri)
    Alpine ibex (Capra ibex)
    Spanish ibex (Capra pyrenaica)
    Siberian ibex (Capra sibirica)
    West Caucasian (Capra caucasica)
    East Caucasian ibex (Capra cylindricornis)
    Nubian ibex (Capra nubiana)
    Walia ibex (Capra walie)



    The evidence for the splits of Groves & Grubbs

    I have devided all splits made by G&G in four categories: Possibly correct, Oversplit, but split possibly correct, Split might be warranted and Split not warranted. In each category I have further indicated for which splits there is enough data and for which splits more research is certainly needed.

    1. Possibly correct

    1.1 Evidence present
    Waterbuck (Kobus ellipsiprymnus)
    Grant's gazelle (Nanger granti)
    Hartebeest (Alcelaphus buselaphus)

    1.2 More research needed

    Bay duiker (Cephalophus dorsalis)
    Chinese goral (Naemorhedus griseus)


    2. Oversplit, but a split possibly correct

    2.1 Evidence present
    Bushbuck (Tragelaphus scriptus)
    Kirk's dikdik (Madoqua kirkii)
    Tsessebe/Topi/Tiang/Korrigum (Damaliscus lunatus)
    Beisa oryx (Oryx beisa)
    Pyrenean chamois (Rupicapra pyrenaica)
    Urial/Mouflon (Ovis vignei)
    African buffalo (Syncerus caffer)

    2.2 More research needed:
    Greater kudu (Tragelaphus strepsiceros
    Klipspringer (Oreotragus oreotragus)
    Suni (Nesotragus moschatus)


    3. Split might be warranted, more evidence is needed

    Mountain reedbuck (Redunca fulvorufula)
    Salt's dikdik (Madoqua saltiana)
    Guenther's dikdik (Madoqua guentheri)
    Blue duiker (Philantomba monticola)
    Weyn's duiker (Cephalophus weynsi)
    Black-fronted duiker (Cephalophus nigrifrons)
    Sumatran goral s.l. (Capricornis sumatraensis)


    4. Split not warranted

    4.1 Evidence present

    Lesser kudu (Tragelaphus imberbis)
    Common reedbuck (Redunca arundinum)
    Oribi (Ourebia oribi)
    Dorcas gazelle (Gazella dorcas)
    Goitered gazelle (Gazella subgutturosa)
    Arabian gazelle (Gazella arabica)
    Indian gazelle (Gazella bennetti)
    Thomson's gazelle (Eudorcas thomsoni)
    Springbok (Antidorcas marsupialis)
    Sharpe's grysbok (Ramphicerus sharpei)
    Impala (Aepyceros melampus)
    Bontebok/Blesbok (Damaliscus pygargus)
    Common wildebeest (Connochaetes taurinus)
    Sable antelope (Hippotragus niger)
    Yellow-backed duiker (Cephalophus silvicultor)
    White-bellied duiker (Cephalophus leucogaster)
    Saiga (Saiga tartarica)
    Alpine chamois (Rupicapra rupicapra)
    Himalayan goral (Naemorhedus goral)
    Argali (Ovis ammon)

    4.2 More research needed
    Sitatunga (Tragelaphus spekii)
    Bohor reedbuck (Redunca redunca)
    Kob (Kobus kob)
    Southern lechwe (Kobus leche)
    Gerenuk (Litocranius walleri)
    Common duiker (Sylvicapra grimmia)
    Ogilby's duiker (Cephalophus ogilbyi)
    Takin (Budorcas taxicolor)
    Wisent (Bos bonasus)



    Additionally there are several species that might need to be lumped based on genetic (and morphological) evidence:
    Natal red duiker & Harvey's duiiker
    Ogilby's duiker & Peter's duiker
    Slender-horned gazelle, Cuvier's gazelle & Sand gazelle
     
    Last edited: 6 Jan 2019
  3. TheGerenuk

    TheGerenuk Well-Known Member 5+ year member

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    Will there be more ungulate groups done?
     
  4. Jurek7

    Jurek7 Well-Known Member 15+ year member

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    Thank Lintworm, very interesting.

    So most of the proposed splits don't stand.

    I notice that you are already applying very easy criteria to split the species anyway. Size difference in Wisent is used as an argument, but bigger size difference in African Buffalo signifies nothing but ecomorphs. Size of white patches is an argument to split Common and Pyrenean chamois, although color patches on wild sheep signify only subspecies.

    It looks like an argument often used is that two forms show some difference consistently (even not so big one). This will however easily lead to error with small sample size or a local inbred sub-population.
     
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  5. lintworm

    lintworm Well-Known Member 15+ year member

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    All the other Ungulate groups will be covered in due time, but it will take some time before this list will be completed.

    As you will have seen, all the splits you mention here are not only based on morphological data, but also on genetic data (see the references at the end of each profile). I wouldn't split Wisent, for a lack of genetic evidence, even though morphologically the animals are quite distinct. We don't know anything about their evolutionary history though. In the case of African buffalo there are size differences between the forest and the savanna buffaloes. The genetic data do however show that the forest ecotype has developed multiple times independently out of a savanna ancestor and all are nested within savanna types. So it is wrong to recognize the forest ones as a different species, as then you would end up with 10+ different buffalo species, which hardly differ genetically.

    In the case of Common and Pyrenean chamois, the morphological difference is backed up with very clear genetic data showing a long time of divergence between two species, this is now partly visible in different color patters, mating behavior as well as in some skull characteristics. Pelage differences between different Urial or Argali subspecies are not backed up with genetic data, as in they have diverged only very very recently and many of the proposed subspecies aren't even monophyletic based on genetic data.

    I have always tried to use a combination of data sources, where possible multiple morphological traits as well as genetic data, most often only Mtdna is available, but where possible also nuclear dna.

    That is indeed the whole problem of the strict PSC approach that G&G have used in their book. Combined with statistical abuse this has resulted in a large number of splits that don't stand, especially when they are integrated with other data sources. G&G are very much old school taxonomists and their work has been really valuable, but with their strict PSC approach they missed the whole story that genetic data can tell.
     
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  6. lintworm

    lintworm Well-Known Member 15+ year member

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    MOSCHIDAE

    The Musk deer family consists of only one genera: Moschus in which 7 species are currently recognized. Though they are called deer, they are actually the closest relatives of the Bovids. There has been considerable taxonomic debate about the number of valid species of Musk deer, ranging from one to seven depending on the author. Currently 7 species are generally recognized, though there still is some debate on the validity of some species and genetic data are mostly lacking.

    Siberian musk deer
    The Siberian musk deer (Moschus moschiferus) is the most widespread musk deer species and occurs from extreme NE Kazakhstan throughout much of E Russia, N Mongolia, N China and the Korean Peninsula. Five subspecies are currently recognized, but their validity is questionable:

    M.m. moschiferus Kazakhstan, Siberia & Mongolia
    M.m turovi Far Eastern Russia
    M.m. arcticus Veryokhansk ridge
    M.m. sachalinensis Sakhalin Island
    M.m. parvipes Korean peninsula

    Moschus moschiferus moschiferus
    [​IMG]

    @Dormitator , Edinburgh Zoo, UK

    [​IMG]
    @Tomek , Zoo Berlin, Germany

    Anhui musk deer
    The Anhui musk deer (Moschus anhuiensis) is a monotypic species that occurs only in the Dabie Mountains in the west of the Anhui Province, China. There has been debate as to whether this species is actually a subspecies of Forest musk deer (Moschus berezovskii)

    No pictures of this species have been uploaded to the gallery.

    Forest musk deer
    The Forest musk deer or Dwarf musk deer (Moschus berezovskii) occurs throughout China and also in N Vietnam. Currently four subspecies are recognized:

    M.b. berezovskii Sichuan, Qinghai, Tibet
    M.b. bjiangensis northwestern Yunnan
    M.b. caobangis Yunnan, Guangxi, Guangdong, northern Vietnam
    M.b. yunguiensis Yunnan-Guizhou highland, Hunan, Jiangxi

    Only one picture of this species has been uploaded to the gallery:
    [​IMG]
    @Deer Forest , Museum of Biological Specimens, Beijing, China

    Black musk deer
    The Black musk deer (Moschus fuscus) is a monotypic species that occurs in the Himalayas in China, Nepal, Bhutan, India and extreme NW Myanmar.

    No pictures of this species have been uploaded to the gallery.

    Alpine musk deer
    The Alpine musk deer (Moschus chrysogaster) is a Musk deer from Central and Southern China, and extreme N Nepal, Bhutan and India. Two subspecies are generally recognized:

    M.c. chrysogaster SE and S Tibet, Nepal, Bhutan, India
    M.c. sifanicus Qinghai, Gansu, Ningxia, western Sichuan and northwestern Yunnan

    No pictures of this species have been uploaded to the gallery.

    Himalayan musk deer
    The Himalayan musk deer (Moschus leucogaster) was formerly considered a subspecies of the Alpine musk deer, but is now generally recognized as a distinct monotypic species. It occurs in the foothills of the Himalayas from Bhutan, through N India, Nepal and extreme S China.

    No pictures of this species have been uploaded to the gallery.

    Kashmir musk deer
    The Kashmir musk deer (Moschus cupreus) was described as a subspecies of Alpine musk deer, but is now generally regarded as a separate monotypic species. This species occurs in the Himalayas in extreme NW India and N Pakistan, the Kashmir Region and N Afghanistan.

    No pictures of this species have been uploaded to the gallery.
     
  7. Chlidonias

    Chlidonias Moderator Staff Member 15+ year member

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    I have put a couple of photos of Ovis ammon darwini in the Mongolia wildlife gallery.
     
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  8. lintworm

    lintworm Well-Known Member 15+ year member

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    CERVIDAE

    Cervidae, the Deer family, is the second largest ungulate family and traditionally +- 55 species of Deer have been recognized, this number was increased somewhat by G&G, but not as crazily as for Bovids. Deer taxonomy on both the species and the genus level is still a hot topic and future changes are to be expected.

    Cervinae

    Muntiacini

    Tufted deer


    The Tufted deer (Elaphodus cephalophus) is a small deer species from China and possibly NE Myanmar. Traditionally four species have been recognized, though G&G do not name any in their book because of lacking data, though the few samples available seem to show large variability.

    E.c. cephalophus SW China, NW Myanmar
    E.c. michianus SE China
    E.c. ichangensis C & S China
    E.c. fociensis distribution unclear, validity doubtful

    E.c. cephalophus
    [​IMG]

    @Maguari, Lowry Park Zoo, Tampa

    E.c. michianus
    [​IMG]

    @Tim May , Wroclaw Zoo, Poland
     
  9. lintworm

    lintworm Well-Known Member 15+ year member

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    Red muntjac species complex

    The Red or Indian muntjac (Muntiacus muntjak) was previously considered as a single species ranging from India to Indonesia. Mainly based on a different chromosome number this species has been split in 2: Northern red muntjac (Muntiacus vaginalis) that ranges from the Indian peninsula and Indochina to N Malaysia and the Southern red muntjac (Muntiacus muntjak) in Malaysia and Indonesia. This split is accepted by e.g. the IUCN but is not free of discussion, mainly because chromosome numbers have only been sampled from very few individuals. In the book Ungulate Taxonomy G&G further split Red muntjak additionally recognizing Muntiacus malabaricus from Sri Lanka and W Ghats, India; Muntiacus aureus NW & C India and parts of Myanmar; Muntiacus nigripes from Hainan, Yunnan and N Vietnam. Muntiacus montanus from the Sumatran highlands.

    Apart from Muntiacus montanus these splits are generally not accepted and the IUCN red list authors can tell exactly why:
    Three species are for now recognized:

    Northern red muntjac
    The Northern red muntjac (Muntiacus vaginalis) ranges from in the Indian Peninsula through Indochina S to N Malaysia at the Isthmus of Kra. Chromosome count 2n=8 in females, 2n=9 in males. The following 8 subspecies have traditionally been recognized, though their validity is sometimes doubtful.

    M.v. vaginalis Nepal. Bhutan, NE India, Bangladesh
    M.v. annamensis S Laos, S Vietnam & Cambodia
    M.v. aureus Pakistan, N & C India
    M.v. curvostylis Myanmar, Thailand
    M.v. malabaricus S India & Sri Lanka
    M.v. menglalis S China, possibly N Indochina
    M.v. nigripes Hainan Peninsula
    M.v. yunnanensis S China

    G&G merge annamensis, menglalis into curvostylis and yunnanensis in nigripes.

    No subspecies:
    [​IMG]
    @Giant Eland , Shadow Nursery, USA

    Muntiacus vaginalis vaginalis
    [​IMG]

    @Chlidonias , Assam State Zoo, India

    Muntiacus vaginalis curvostylis
    [​IMG]
    @Giant Eland, Khao Keow Forest & Wildlife Reserve Park, Thailand

    Southern Red muntjac

    The Southern red muntjac (Muntiacus muntjak) is a monotypic species from Malaysia, S of the Isthmus of Kra and Sumatra, Java, Borneo, Bali and associated islands. Chromosome count 2n=6 in females, 2n=7 in males.

    [​IMG]
    @Giant Eland , Taman Safari Bogor, Indonesia

    Sumatran mountain muntjac

    The Sumatran mountain muntjac (Muntiacus montanus) is a species of questionable validity of which hardly anything is known. This species is limited to mountanous areas of W Sumatra, Indonesia.

    No pictures of this species have been uploaded to the gallery.
     
  10. Chlidonias

    Chlidonias Moderator Staff Member 15+ year member

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    I didn't know they had been split. I'll have to have a look at this and see what I think of it - I might be able to add an armchair tick!
     
  11. Chlidonias

    Chlidonias Moderator Staff Member 15+ year member

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    Turns out I don't think much of that split. It is likely I actually already knew about it and forgot again because (to quote the IUCN) it "rests on little evidence". Phrases used throughout the IUCN taxonomy account are in the vein of "postulated", "assumption", "assumed", "purported", "minor", and it ends with "There appears to have been no significant further investigation of this taxonomic split since 2008, and thus the 2014 reassessment largely for the sake of status quo maintains this taxonomic treatment".


    Groves (2003) elected to raise non-Sundaic forms of M. muntjak (s.l.) from subspecific taxa to the species M. vaginalis, leaving the Sundaic forms to constitute M. muntjak (s.s.). In 2008 the IUCN Red List chose to follow this split, with the following caveats. This position, already postulated by previous authors (e.g. Groves and Grubb 1990), rests on little evidence, primarily the assumption that all taxa within M. muntjak in the Sunda region carry a unique karyotype different from all M. vaginalis in northern regions. However, the Sundaic karyotype seems to have been documented in only a few individuals from the Malay Peninsula south of the Isthmus of Kra (Wurster and Aitkin 1972, Groves and Grubb 1990, Groves 2003, Tanomtong et al. 2005), assumed to be M. muntjak (the type locality of the later is Java). Other purported differences (e.g. dorsal darkening and short nasals, Groves 2003, Groves and Grubb 1990), if they can be considered diagnostic characters rather than traits (see below), appear minor rather than ones likely to separate species-level taxa. Although differences in chromosome number between muntjacs appears likely to constitute a good species boundary, preventing extensive interbreeding (see Groves and Grubb 1990), a much wider sampling of karyotype (especially within the presumed range of M. muntjak (s.s.)) is needed to place on solid ground the systematic position which assumes separate and widespread ranges for the two so far identified karyotypes. There appears to have been no significant further investigation of this taxonomic split since 2008, and thus the 2014 reassessment largely for the sake of status quo maintains this taxonomic treatment.
     
  12. TheGerenuk

    TheGerenuk Well-Known Member 5+ year member

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    Glad to see that this thread is still active! Keep up the good work!
     
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  13. ThylacineAlive

    ThylacineAlive Well-Known Member 10+ year member

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    The US population has traditionally listed as Muntiacus muntjak vaginalis. Would this not translate into M. v. vaginalis once split, or were the subspecies not described until after the split?

    ~Thylo
     
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  14. Chlidonias

    Chlidonias Moderator Staff Member 15+ year member

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    The subspecies were already in place - they have just split the species in two (and apparently dropped any subspecies which were formerly recognised in the Sunda region, to make muntjak monotypic).
     
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  15. UngulateNerd92

    UngulateNerd92 Well-Known Member Premium Member 5+ year member

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    I am sorry to bump this thread, but I can't help but to wonder what Colin Groves' and Peter Grubb's rebuttals would be to the findings/analysis done by @lintworm here, if they were still alive.

    Great work by the way! I can't thank you enough for putting this thread together!
     
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  16. lintworm

    lintworm Well-Known Member 15+ year member

    Joined:
    27 Oct 2008
    Posts:
    5,498
    Location:
    Europe
    Hog deer

    The Hog deer (Axis porcinus) is a small species of deer from Southeast Asia. This species has recently been placed in the genus Hyelaphus and one genetic study placed it as a sister to Rusa instead of Axis deer, but later genetic studies have confirmed that this was a mistake and Hog deer are indeed sister species to Axis deer and do not warrant recognition of a separate genus as the estimate date of divergence is only 2.6 MYA. Traditionally two subspecies have been recognized:

    A.p. porcinus Indian hog deer; Pakistan, N and NE India, Bangladesh and Nepal
    A.p. annamiticus Indochina hog deer; Myanmar, Cambodia & Manipur, India

    The exact historic distribution is unclear and only recently was a population in E India reliably assigned to annamiticus. Both subspecies are elevated to species status by G&G

    Sample sizes

    Skins
    No sample sizes are given

    Skulls
    porcinus 5-6
    annamiticus 2-4

    Skins

    annamiticus is described as being much brighter, more ochery and less grey and buffy compared to porcinus. annamiticus is also not speckled.

    Skulls

    porcinus and annamiticus cannot be distinguished based on skull characteristics.

    Additional data

    There has been one recent study that compared porcinus and annamiticus hog deer genetically (Gupta et al. 2018). This study found a sequence divergence of +- 4% and has an estimated time of divergence of 220.000 years ago (95% HPD 110.000-338.000 years ago).
    From personal observations in zoos it seems that there is high variation in coat colouration between porcinus Hog deer between individuals and throughout the year, with colour ranging from ochery to dark grey, which is used here to differentiate between annamiticus and porcinus, the more ochery coloured porcinus do however normally have spots.

    Summarizing

    The genetic data indicate the existence of two clearly differentiated subspecies that might be approaching species status, though on PSC criteria they represent clearly separate species. Morphological differences are however very slight, which could be due to the extremely low sample sizes used by G&G. More information on the exact distribution and improved morphological analyses, including antlers, would be necessary to assess whether annamiticus indeeds should have species status. For now treating them as a separate management unit is certainly necessary.

    Axis porcinus porcinus

    Male
    [​IMG]
    @Goura Burgers' Zoo, Arnhem, Netherlands

    Female
    [​IMG]
    @Chlidonias Kaziranga NP, India

    Axis porcinus annamiticus
    [​IMG]

    @Giant Eland , Khao Kheow Forest & Wildlife Reserve Park, Thailand

    Reference

    Gupta et al. (2018): Genetic analysis of endangered hog deer (<i>Axis porcinus</i>) reveals two distinct lineages from the Indian subcontinent
     
    Last edited: 13 May 2019
  17. lintworm

    lintworm Well-Known Member 15+ year member

    Joined:
    27 Oct 2008
    Posts:
    5,498
    Location:
    Europe
    Apart from Hog deer, three other species are recognized in the genus Axis

    Axis deer

    The Axis deer (Axis axis) is a medium-sized deer from India, Sri Lanka, Nepal, Bhutan and Bangladesh. It is considered monotypic

    Male
    [​IMG]
    @Chlidonias , Wilpattu NP, Sri Lanka

    Female
    [​IMG]
    @devilfish , Tatu Carreta, Argentina

    Bawean deer

    The Bawean deer (Axis kuhlii) is a monotypic critically endangered deer species that is restricted to the island of Bawean between Java and Borneo.

    Male
    [​IMG]
    @Maguari , Edinburgh Zoo, UK

    Female
    [​IMG]
    @Giant Eland , Taman Safari Bogor, Indonesia

    Calamian deer

    The Calamian deer (Axis calamianensis) is a monotypic critically endangered deer that is restricted to the Calamian Islands in the Phillipines.

    Male
    [​IMG]
    @Blackduiker , Los Angeles Zoo, USA

    Male & female
    [​IMG]
    @Ituri , Phoenix Zoo, USA
     
  18. lintworm

    lintworm Well-Known Member 15+ year member

    Joined:
    27 Oct 2008
    Posts:
    5,498
    Location:
    Europe
    The sister genus to Axis is Rucervus, until recently Brow-antlered deer were also assigned to this genus, but multiple studies that worked with Mtdna have clearly shown that Brow-antlered deer are sister to Pere-david deer and is sister to Cervus. This means that either Cervus will need to be enlarged or Brow-antlered deer should be placed in the monotypic genus Panolia.

    The genus Rucervus now consists of only one extant and one extinct species.

    Barasingha
    The Barasingha or Swamp deer (Rucervus duvaucelii) is the only extant member of the genus Rucervus. This is a large deer from India and S Nepal and three subspecies are generally recognized:

    R.d. duvaucelii N India, S Nepal
    R.d. branderi C India
    R.d. ranjitsinhi NE India

    G&G elevate all subspecies to species status.

    Sample sizes
    No new data seem to have been used and all shown results seem to be based on Groves (1983).

    Skins
    No sample sizes are given

    Skulls/antlers
    duvaucelii 9-13
    ranjitsinhi 2-3
    branderi 2-5

    Sample sizes are thus mostly extremely limited.

    Skins
    No details on skins are given for branderi except that this species has "well-knit" feet with hairy pasterns, possibly an adaptation to living in less marshy areas than the other Barasingha. ranjitsinhi is described as having less prominent white spots in moult, short tail and small pointed ears with very little white hair on the inside when compared to duvaucelii.

    Skulls/antlers
    Several differences in antlers and skulls are described by G&G. Though these might be true when looking at the averages, there is however wide overlap in these measurements when looking at the standard deviation. Taking into account the extremely small sample sizes for especially the skull measurements, this means no conclusion can be drawn from these data. The only possible difference might be that branderi is relatively small with relatively large antlers, but given the paucity of data this cannot be reliably concluded.

    Additional data
    A recent genetic study (Kumar et al. 2017) using both nuclear and Mtdna found that each subspecies is genetically different. They did however find evidence for relatively recent colonization, though the date of divergence was not estimated. The genetic distance between the taxa was relatively high and would approach species status. Given the fact that genetic distance increases faster in small isolated populations, which was likely the case for Barasingha, this might however not equal species status.

    Summarizing
    Overall there seems to be no strong case for splitting Barasingha, except if you strongly favour a PSC approach. Given the current morphological data, differences can hardly be identified reliably. The genetic results show that the three subspecies are clearly identifiable, but differences are not huge and estimated times of divergence are not known. Future research on Barasingha should however be a priority to assess how long the different populations have been isolated and whether (and which) morphological differences do exist.

    It remains unclear to which subspecies captive Barasingha in Europe and America belong, though they are often labelled as duvaucelii. The only Barasingha of which pictures have been uploaded from the wild are from Kaziranga, India.

    Rucervus duvaucelii ranjitsinhi
    [​IMG]

    @Chlidonias , Kaziranga NP, India

    Zoo Barasingha; male
    [​IMG]
    @ro6ca66 , ZSL Whipsnade Zoo, UK

    Female
    [​IMG]
    @ro6ca66 , ZSL Whipsnade Zoo, UK

    Schomburgk's deer

    The Schomburgk's deer (Rucervus schomburgki) is an extinct deer species that originally occurred in Thailand, though rumours of continued survival exist, this species is generally considered to be extinct since 1938.

    [​IMG]
    Uploaded by @Joker1706 , Zoo Berlin, Germany


    References

    Groves (1983): https://www.biodiversitylibrary.org/page/48745060#page/686/mode/1up

    Kumar et al. (2017): Mitochondrial and Nuclear DNA Based Genetic Assessment Indicated Distinct Variation and Low Genetic Exchange Among the Three Subspecies of Swamp Deer ( Rucervus duvaucelii)
     
  19. ThylacineAlive

    ThylacineAlive Well-Known Member 10+ year member

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    20 Oct 2012
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    10,699
    Location:
    Connecticut, U.S.A.
    What about the small number of branderi listed for some European zoos? Most are listed as nominate, but some of the French collections and a single animal at Port Lympne are listed as branderi.

    ~Thylo
     
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  20. lintworm

    lintworm Well-Known Member 15+ year member

    Joined:
    27 Oct 2008
    Posts:
    5,498
    Location:
    Europe
    Zootierliste states that based on the first ISB it is unclear where all the founder animals were sourced and it is thus impossible to assess to which subspecies they belong... I have not been able to find any other information on their origin, except it being stated somewhere that most if not all American & European are likely duvaucelii... But if someone else has new information that would shed light on the status of these branderi specimens, that would be great...