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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. lintworm

    lintworm Well-Known Member 15+ year member

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    ANTILOCAPRIDAE

    The Antilocapridae is the smallest ungulate family and consists of only one extant species.

    Pronghorn

    The Pronghorn (Antilocapra americana) is a unique species of ungulate endemic to North America. Originally up to five subspecies were recognized, but oregona and mexicana are now largely considered invalid. There is also debate as to whether the three remaining subspecies are valid, or whether this species should be considered monotypic. Here three subspecies are provisionally recognized:

    A.a. americana from S Canada to N and C Mexico
    A.a. peninsularis N Baia California, Mexico
    A.a. sonoriensis, S Arizona, USA & N Sonora, Mexico.

    A.a. americana
    [​IMG]

    @Giant Eland , Bryce Canyon NP, USA

    A.a. peninsularis
    [​IMG]
    @ThylacineAlive , Los Angeles Zoo, USA

    [​IMG]
    @Blackduiker , Los Angeles Zoo, USA
     
    Last edited: 10 Nov 2019
  2. ThylacineAlive

    ThylacineAlive Well-Known Member 10+ year member

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    While I am far from an expert on the subject, my understanding is that at least americana and peninsularis are both rather reliant on the habitat they're found in. For instance, Cougars from anywhere in North America do just fine in "Florida Panther" habitat. With Pronghorn, however, you could not put a peninsularis in the range of americana and expect it to survive. In general, americana have proven to be rather difficult to keep in captivity outside of their native range as well. This is why the AZA peninsularis program is entirely restricted to the arid areas of the Southwest and why all Pronghorn populations outside of their native range seem to be slowly disappearing. I'm not sure if this is observable in sonoriensis as well but I would expect so.

    ~Thylo
     
  3. lintworm

    lintworm Well-Known Member 15+ year member

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    TRAGULIDAE

    The Chevrotain or Mouse deer family (Tragulidae) is distributed mostly in SE Asia, with one species occuring in Africa. There is considerable debate with regards to how many species should be recognized, but most sources provisionally list 10 species.

    Water chevrotain

    The Water chevrotain (Hyemoschus aquaticus) is the largest Chevrotain species and the only one occuring outside of Asia. It is currently distributed throughout most of West and Central Africa, from Guinea in the west to W Uganda in the east. The most southern record stems from N Angola. Three subspecies have been named (aquaticus, batesi, cottoni) but their validity is questionable.

    [​IMG]
    @Tim May , Natural History Museum Brussel, Belgium


    White-spotted chevrotain

    The White-spotted chevrotain (Moschiola meminna) is one of currently three recognized species in the genus Moschiola. Groves & Meijaard (2005) split the genus Moschiola in three species based on morphological analysis. They found clear differences in skull size and in color pattern between populations from India (indica) and between the Sri Lankan dry zone (meminna) and wet zone (kathygre) populations. Even though the sample size in this study was limited and no genetic data backing up this split are available, this split has been provisionally accepted by most recent authors. No new genetic evidence has been presented for this three-way split yet and this should be considered a high priority. Zurano et al. (2019) in their phylogeny for all Cetartiodactyla based on complete mitochondrial genomes included 1 sample of meminna and one of indica and their estimates give a time of divergence of roughly 5 million years ago, which is an indication that species status might be deserved.

    No pictures of this species have been uploaded to the gallery

    Yellow-striped chevrotain

    The Yellow-striped chevrotain (Moschiola kathygre) is the smallest Moschiola species and has in the past been treated as conspecific with Moschiola meminna. It is provisionally recognized as a separate species based on Groves & Meijaard (2005). This species occurs in the wet zone of Sri Lanka

    [​IMG]
    @baboon , Singapore Zoo, Singapore

    [​IMG]
    @Chlidonias , National Zoological Gardens of Sri Lanka, Colombo, Sri Lanka

    Indian chevrotain

    The Indian chevrotain (Moschiola indica) was previously considered conspecific with the other Moschiola species, but is provisionally recognized as a monotypic species here based on Groves & Meijaard (2005). This species occurs throughout a large part of the Indian subcontinent and might be present / have been present in Bangladesh and Nepal.

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

    Greater Indo-Malayan chevrotain

    The Greater Indo-Malayan chevrotain, or Greater chevrotain (Tragulus napu) is a large species of Chevrotain that occurs in S Myanmar, S Thailand, Malaysia, Singapore (Pulau Ubin), Borneo, Sumatra and adjacent islands. A large number of subspecies has been described but seven are currently recognized:

    T.n. napu S Myanmar, Malay Peninsula, Borneo, S Sumatra
    T.n. banguei Banggi and Balenbangan Islands, N off Borneo
    T.n. bunguranensis Natuna Island off W Borneo
    T.n. neubronneri, N Sumatra
    T.n. niasis, Nias Island off W Sumatra
    T.n. rufulus Tioman I, Riau and Lingga Archipelagos
    T.n. terutus, Terutau I of W Malay Pensinsula

    Tragulus napu napu
    [​IMG]

    @Chlidonias , Zoo Melaka, Malaysia


    [​IMG]
    @Ding Lingwei , Bronx Zoo, New York, USA

    I have seen the Chevrotain at Singapore Zoo been referred to as subspecies rufulus, but not pictures of these animals seem to have been uploaded.

    Balabac chevrotain

    The Balabac chevrotain (Tragulus nigricans) was formerly considered a subspecies of the Greater Indo-Malayan chevrotain, but has been upgraded to species status as it is morphologically very distinct. This species is restricted to Balabac, Bugsuk & Ramos Islands in the Palawan region of the Philippines.

    [​IMG]
    @vogelcommando , Diergaarde Blijdorp, Rotterdam, Netherlands

    Silver-backed chevrotain

    The Silver-backed chevrotain (Tragulus versicolor) is a species of Chevrotain that is restricted to SE Vietnam and possibly Laos and Cambodia, data on this species are very rare, also because it likely overlaps with T. kanchil and a lack of surveys.

    No picture of this species has been uploaded to the gallery.

    Lesser Indo-Malayan chevrotain

    The Lesser Indo-Malayan chevrotain (Tragulus kanchil) is a small Chevrotain occuring in most of mainland SE Asia, Singapore, Sumatra, Borneo and adjacent islands. Initially animals from Java were included in this species, but they were split by Meijaard & Groves (2004) based on morphological analysis. 16 subspecies are provisionally recognized, but this is likely an overestimate. Genetic research by Endo et al. (2004) found three clusters within Lesser Indo-Malayan chevrotain, animals from Borneo formed a separate cluster from mainland animals, where two groups were also identified, one north and the other south from the Isthmus of Kra. No animals from Sumatra and Java were sampled. Meijaard & Groves (2004) already commented animals from Borneo might represent a separate species and this seems to be backed up by genetic data. New genetic research, including Chevrotain from over the whole range, would be necessary to shed light on the Lesser chevrotain of SE Asia and all Indonesian islands. Meijaard & Groves (2004) recognize the following 16 subspecies, but a revision is necessary:

    T.k. kanchil, Sumatra and Mendol & Berhana Islands of Sumatra
    T.k. abruptus, Subi Island off Borneo
    T.k. affinis, Vietnam, Laos SE & E Thailand
    T.k. anambensis, Anambas Archipelago
    T.k. angustiae, S Myanmar, SW Thailand,
    T.k. everetti, Natuna Island off Borneo
    T.k. fulviventer S Malay Peninsula
    T.k. hosei, Borneo
    T.k. klossi, N Borneo
    T.k. luteicollis, Bangka Island off Sumatra
    T.k. pidonis, Koh Pipidon Island of Malay Peninsula
    T.k. ravulus, islands of W Malay Pensinsula
    T.k. ravus, S Thailand, N Malaysia
    T.k. rubeus, Riau Archipelago
    T.k. siantanicus, Anambas Archipelago

    Animals in European zoos are mostly signed as Tragulus javanicus, but are most likely Tragulus kanchil or hybrids. Many animals are related to an import by Poznan Zoo of Tragulus kanchil affinis, but it is unknown what the other import locations were and if multiple species were mixed (see The mousedeer headache). Subspecies of zoo animals are mostly based on this link as well.

    Tragulus kanchil affinis
    [​IMG]

    @Giant Eland , Dusit Zoo, Thailand

    Tragulus kanchil angustiae
    [​IMG]

    @Chlidonias , Yadanabon Zoo, Myanmar

    Tragulus kanchil fulviventer
    [​IMG]
    @Chlidonias , Taman Negara NP, Malaysia

    Tragulus kanchil ravus
    [​IMG]

    @Chlidonias , Penang Bird Park, Malaysia

    Tragulus kanchil klossi
    [​IMG]

    @lintworm , Singapore Zoo, Singapore

    Tragulus kanchil (X javanicus?)
    [​IMG]

    @ro6ca66 , Paignton Zoo, UK,

    Javan chevrotain

    The Javan chevrotain (Tragulus javanicus) was formerly considered conspecific with Tragulus kanchil, but has been split on morphological grounds. There is no good genetic research backing up this split, but Zurano et al. (2019) estimated based on the mitochondrial genome of 1 T. kanchil and 1 T. javanicus that the time of divergence was roughly 2.5 mya.

    [​IMG]
    @Chlidonias , Taman Mini Indonesia, Jakarta, Indonesia

    Northern chevrotain

    The Northern chevrotain (Tragulus williamsoni) was formerly considered conspecific with Tragulus kanchil, but has been elevated to species status by Meijaard & Groves (2004). There is no genetic research to back this up, but a somewhat extended dataset seems to confirm this placement (Meijaard et al., 2017). Genetic research would however still be a priority.

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


    References

    Groves & Meijaard (2005): https://pdfs.semanticscholar.org/adf2/07360161a2da1fdfa71e8fecfd09ae448c82.pdf

    Meijaard & Groves (2004): http://doc.rero.ch/record/210302/files/PAL_E4353.pdf

    Meijaard et al. (2017): https://www.researchgate.net/profil...e-least-documented-mammal-species-in-Asia.pdf

    Zurano et al. (2019): http://www.chufpb.com.br/danmesq/Publicacoes_files/1-s2.0-S1055790318302720-main.pdf
     
  4. lintworm

    lintworm Well-Known Member 15+ year member

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    HIPPOPOTAMIDAE

    The Hippopotamus family consists of two genera, each with one extant species. Until recently 2 now extinct Hippopotamus species lived on Madagascar, which might have survived until modern times in relict populations. G&G additionally recognize the now extinct Niger Delta pygmy hippopotamus as a distinct species, rather than as a subspecies.

    Common hippopotamus

    The Common hippopotamus (Hippopotamus amphibius) is a large mammal, endemic to Africa. This species is distributed patchily throughout sub-saharan Africa and is now extinct in Algeria, Egypt, Eritrea, Mauritania and Liberia. Three subspecies are generally recognized and this is backed up by small genetic differences between them (Okello et al., 2005).

    H.a. amphibius, most of sub-saharan Africa
    H.a. capensis, Zambia, south to S Africa
    H.a. kiboko, Kenya & Somalia


    Hippopotamus amphibius amphibius
    [​IMG]

    @lintworm , Katavi NP, Tanzania

    Hippopotamus amphibius capensis
    [​IMG]

    @Maguari , Khwai Community Area, Botswana

    Hippopotamus amphibius kiboko
    [​IMG]

    @LaughingDove , Lake Naivasha, Kenya

    Zoo animals:
    [​IMG]
    @Arek , Zoo Wroclaw, Poland

    [​IMG]
    @Arek , Zoo Wroclaw, Poland

    Pygmy hippopotamus

    The Pygmy hippopotamus (Choroepsis liberiensis) is a small hippo species that occurs from Guinea to Ivory Coast, with an extinct population in the Niger delta in Nigeria. Traditionally two subspecies have been recognized:

    C.l. liberiensis Guinea to Ivory Coast
    C.l. heslopi Niger Delta to Cross River, Nigeria (extinct)

    G&G elevate heslopi to species status based on measurements by Corbett (1969). Even though the sample sizes used are extremely limited (3 for heslopii and 3-14 for liberiensis) the few heslopi skulls are distinct in many measurements from liberiensis skulls. There is no genetic data to back this up, as heslopi has not been seen since 1945, but the limited dataset does show that it is very possible heslopi was a separate species and not a subspecies. Both taxa are separated by the Dahomey gap and there are multiple examples of species separated by this gap. It is impossible to assess the true status of heslopi, but from a PSC perspective this is a clear case of heslopi being a separate species.

    Choroepsis liberiensis liberiensis
    [​IMG]

    @m30t , Toronto Zoo, Canada


    References

    Corbett (1969): ZSL Publications

    Okello et al. (2005): Mitochondrial DNA variation of the common hippopotamus: evidence for a recent population expansion
     
  5. Arizona Docent

    Arizona Docent Well-Known Member 15+ year member

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    @lintworm Interesting note about pronghorns - I had not heard that A.a. mexicana (which is what my photo represents under the old taxonomy) has been lumped with A.a. sonoriensis. Although I admit from a visual standpoint all pronghorns look the same to me.
     
  6. lintworm

    lintworm Well-Known Member 15+ year member

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    Sorry, my mistake, I didn't look at the map correctly and just noted Sonoita was in Arizona, but it is actually outside the range of sonoriensis, so the picture shows the nominate subspecies, which is what mexicana was lumped with.
     
  7. Chlidonias

    Chlidonias Moderator Staff Member 15+ year member

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    IUCN says this "species" is known from only one specimen - is that still accurate?
     
  8. lintworm

    lintworm Well-Known Member 15+ year member

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    The count is now 2, with a second specimen from Yunnan, China., both specimens fall outside the range of measurements of other Lesser chevrotain. But it is still a taxon that is recognized somewhat preliminary. See: https://www.researchgate.net/public...f_the_least-documented_mammal_species_in_Asia for the latest info on this putative species.
     
  9. lintworm

    lintworm Well-Known Member 15+ year member

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    TAYASSUIDAE

    The Peccary family consists of three genera, in which traditionally one species each has been recognized.

    Collared peccary

    The Collared peccary (Pecari tajacu) is a widely distributed species occuring from the S of the USA throughout Central America and much of tropical South America. Up to fourteen subspecies have been recognized, but the validity of these is doubtful.

    P.t. tajacu Amazon Basin to N Argentina
    P.t. angulatus S USA through NE Mexico
    P.t. bangsi C Panama along NW Colombia
    P.t. crassus EC Mexico
    P.t. crusnigum Nicaragua to W Panama
    P.t. humeralis WC Mexico
    P.t. nanus SE Mexico (Cozumel Island)
    P.t. nelsoni S Mexico to Belize & Guatemala
    P.t. nigrescens S Guatemala, El Salvador & Honduras
    P.t. patira N South America & Trinidad & Tobago
    P.t. sonoriensis SW USA
    P.t. torvus NW South America
    P.t. yucatanensis SE Mexico to N Guatemala

    G&G recognize three separate species:

    tajacu
    (including patira as subspecies) most of South America
    angulatus (including sonoriensis, humeralis) dry zone of N Mexico and S USA
    crassus (including yucatanensis, nanus, crusnigum, angulatus, nelsoni, nigrescens, bangsi, torvus) humid rain forest zone of S Mexico, Central America into NW Colombia and Ecuador W of Andes.

    Sample size

    No sample sizes are given for skins.

    Skulls
    USA 16
    Sinaloa, Mexico 6
    Veracruz, Mexico 2
    Oaxaca, Tabasco, Guatemala, Belize 8-9
    Colombia "N type" 16-18
    Colombia "S type"2
    E Peru, E Ecuador 10-14
    C Peru 5
    Trinidad 1
    Guyana, Suriname, Roraima, 14
    Para 5
    Amazonas 1-6
    Mato Grosso, E Bolivia 8
    N Argentina, Paraguay 5-8
    Sao Paulo, Santa Catarina 2

    Overall quite a large number of samples are present in the analysis, covering most of the distribution of the species.


    Skins

    No description is given how tajacu skins differ from angulatus and crassus. angulatus skins are described as hair banding blackish and gray or white, head lighter in tone, collar white, but often not marked, sometimes with a black dorsal stripe. crassus skins are described as being coarser, wide light hair bands, so the overall colour becomes gray or white, black dorsal stripe and the collar various, often indistinct.


    Skulls

    There is a remarkable difference between skulls of angulatus + crassus and tajacu. The N type skull has been described as "malar crest slopes forward, ending above the canine alveolus; nasal bones pinched or angulated; first upper premolar quadritubercular, with intermediate tubercles, molariforn; molars wrinkled. The S type skull (tajacu) is described as "malar crest directed upward, ending above the infraorbital foramen; nasal bones more rounded on cross section; first upper premolar premolariform; molars not wrinkled.

    G&G indicate they find little overlap between angulatus and crassus skulls with angulatus skulls said to be larger, broader occiput and narrower palates. The evidence for this seems however very scant based on the measurements they provide and there appears to be large overlap. angulatus is also sait to have a larger and narrower skull then tajacu but there seems to be just as much variation within each group as there is between them. G&G go on to describe variation between regions within S America (tajacu), but evidence for their claims is often not evident from the data.


    Additional data

    Gongora et al. (2006) have analyzed the differences between Collared peccary based on both mitochondrial and nuclear dna. They found evidence for two major clades, which correspond with the N (angulatus + crassus) and S skull types (tajacu). There seems to be a contact zone between these clades in Colombia as hybrids between both clades were found there. The genetic difference between the two clades is as big as between Chacoan and White-lipped peccary, which are generally grouped in two separate genera.

    Additionally a difference in chromosomes has been described between both clades. Although both clades have the same number of chromosomes, chromosomes 1 and 8 have different shapes between both clades.

    van Roosmalen described a new Collared peccary species from the Amazon and called it Pecari maximus, based on a single specimen. Genetic research by Gongora et al. (2011) showed however that this specimen falls within Pecari tajacu and there is not reason to recognize this putative species.


    Summarizing

    Combined evidence of skulls, mtdna, nuclear dna and chromosomes point to the existence of two separate clades of Collared peccary, which likely represent two separate species. It would however still be interesting to study the hybrid zone in Colombia to see whether hybrids have lower fitness or whether they interbreed freely. Based on current evidence two species of Collared peccary seem well supported, but there is no support to split the Northern collared peccary in two species (angulatus & crassus). There also seems no evidence for fourteen separate subspecies, but the genetic work found some minor clades within the N and S group, so there is evidence for the existence of subspecies, but their exact boundaries still need to be determined.


    Northern collared peccary: Pecari (tajacu) angulatus
    [​IMG]

    @Ituri , Phoenix Zoo, USA

    [​IMG]
    @Arizona Docent , Tonto Natural Bridge State Park, USA

    "crassus"
    [​IMG]
    @vogelcommando , Parque Zoologico de Centenario, Mexico

    Southern collared peccary: Pecari (tajacu) tajacu
    [​IMG]
    @devilfish , Santa Cruz Zoo, Bolivia

    [​IMG]
    @carlos55 , Zoo Mendoza, Argentina

    I do not know the origin from the European zoo population.

    References

    Gongora et al. (2006): Phylogenetic divisions among Collared peccaries (Pecari tajacu) detected using mitochondrial and nuclear sequences - ScienceDirect

    Gongora et al. (2011): https://s3.amazonaws.com/academia.e...a7a9ed903e08f152c0f271126d017c9d633071c6ce82b
     
  10. lintworm

    lintworm Well-Known Member 15+ year member

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    White-lipped peccary

    The White-lipped peccary (Tayassu pecari) is a widespread species occuring from S Mexico throughout South America and much of tropical South America. Five subspecies have been provisionally recognized, but there is no genetic work backing up these claims:

    T.p. pecari Colombia, Venezuela, the Guianas, Suriname & N Brazil
    T.p. aequatore SW Colombia & Ecuador
    T.p. albirostre S Brazil, E Peru, Bolivia, Paraguay & N Argentina
    T.p. ringens SE Mexico to Nicaragua
    T.p. spiradens Costa Rica to N Colombia

    G&G recognize 4 White-lipped peccary subspecies, not species, by merging ringens with spiradens. Differences in chromosome form have been found between animals from Costa Rica and between two regions in Brazil, but this has not been linked with subspecific distributions or other genetic work.

    T.p. albirostre
    [​IMG]

    @Maguari , Espace Animalier de la Haute Touche, Obterre, France

    T.p. ringens
    [​IMG]

    @carlos55 , Zoo de San Juan de Aragon, Mexico


    T.p. pecari / aequatore
    [​IMG]

    @Giant Eland , Bioparque Los Ocarros, Colombia

    Chacoan peccary

    The Chacoan peccary (Catagonus wagneri) is the largest Peccary and was previously only known from fossil evidence, but in 1971 it was "discovered" that this species was alive and distributed in the Gran Chaco area of Bolivia, Paraguay and Argentina.

    [​IMG]
    @Tomek , Zoo Wroclaw, Poland
     
  11. TeaLovingDave

    TeaLovingDave Moderator Staff Member 10+ year member

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    I'm given to understand that some of the stock at Faunia in Madrid was imported directly from South America and is nominate - not that this matters for the purposes of this thread, as you already have this taxon illustrated ;)
     
    ThylacineAlive likes this.
  12. ThylacineAlive

    ThylacineAlive Well-Known Member 10+ year member

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    For interests sake, the peccaries at Orange County Zoo are nominate angulatus and the ASDM animals are sonoriensis. After that I do not know the status of American captive peccaries.

    ~Thylo
     
  13. lintworm

    lintworm Well-Known Member 15+ year member

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    Babirusa

    The Babirusa (Babyrousa babyrussa s.l.) is a potential species complex from Sulawesi and a few neighbouring islands. Groves (1980) listed only one species, but a revision, based solely on a change of species concept, elevated the four described subspecies, one of them extinct, to species level (Meijaard & Groves, 2002).

    The subspecies were described as follows:
    B.b. babyrussa Sula Islands and Buru (Moluccas archipelago)
    B.b. celebensis N Sulawesi (with status of animals in most of Sulawesi unclear)
    B.b. togeanensis Togian Archipelago
    B.b. bolabatuensis SW peninsula of Sulawesi, extinct


    Sample size

    All work is based on Groves (1980):

    Skins
    babyrussa 2
    celebensis 12
    togeanensis 11
    bolabatuensis 0

    Skulls (males)
    babyrussa 5-6
    celebensis 16-29
    togeanensis 7-36
    bolabatuensis 1

    Some measurements are also available for females, but those have extremely low sample sizes, so are ignored here. Overall 2 of the taxa are reasonably good representated two others not so much.


    Skins

    babyrussa skins are described as always having long and thick hair, with a well developed tail tuft. togeanensis has less long and dense hair than babyrussa, but also a well developed tail tuft. celebensis hair is short and sparse, with only a small tail tuft.


    Skulls

    babyrussa skulls are somewhat shorter than celebensis and togeanensis, but there is overlap in measurements with celebensis, in other skull characteristics differences are minimal. togeanensis are on average somewhat larger than celebensis, with a smaller toothrow, but especially in the size there is wide overlap. bolabatuensis is described as a small version of togeanensis, but as it is based on only one skull it is hard to assess the validity of this taxon.

    When it comes to dentition togeanensis has somewhat smaller teeth than celebensis, contrasting with the on average somewhat larger skull. When it comes to teeth celebensis and babyrussa are relatively similar when looking at average sizes. Upper canines of togeanensis are described as "usually short, slender, somewhat rotated forward, always converging".babyrussa upper canines are described as: "Upper canines of males usually short, slender, with alveolus forwardly rotated, so that lower canine crosses upper in lateral view; generally divergent or parallel to each other, or weekly convergent". celebensis upper canines are described as follows: "Upper canines of males generally long and thick, the alveoli vertically implanted, so that upper canine emerges vertically and is not crossed by lower in lateral view; converging in almost all cases". Note that the illustration in HMW 2 of togeanensis shows forward pointing upper canines, but see plate 2 in Groves (1980), that this is wrong. The differences in canine shape are exaggerated anyway in that plate, when comparing with pictures of skulls in Groves (1980) and comparison in the latter is difficult as for togeanensis a sub-adult skull is pictured, compared adult skulls.


    Additional data

    There is one major study that has looked at both morphological and genetic differentiation between Babirusa across the whole range (Frantz et al., 2018). They show that based on tooth data babyrussa and celebensis are somewhat similar, babyrussa has on average smaller teeth, but there is overlap with celebensis and in the multivariate analysis it does not group completely apart from celebensis. togeanensis does however have far smaller teeth than the other 2 taxa and there is very little overlap. In a multivariate analysis togeanensis is grouped further away from the other Babirusa taxa than Sulawesi warty pig.

    Genetic data do show that there are six major haplogroups, 1 representing togeanensis, 1 representing babyrussa and 4 across different parts of Sulawesi. The genetic data show that babyrussa is the most distinct and that all three taxa are monophyletic. The genetic distance between the haplogroups is however low with variation between populations only being 27% of the total variation present in the dataset.

    Summarizing

    Although this split is accepted provisionally by the IUCN, the genetic results by Frantz et al. (2018) do not support a split. Genetic differentiation between the proposed species is low and only under a narrow PSC view would that still mean there are multiple Babirusa species. Previous research by Groves did not incorporate data from S, C and E Sulawesi, but there is some genetic structuring across the island that might represent multiple subspecies or multiple ESUs. This has a consequence for captive Babirusa as there exact geographic origin is probably unknown

    Babyrousa babyrussa celebensis
    [​IMG]
    @gentle lemur , Chester Zoo, UK

    [​IMG]
    @Giant Eland , Ragunan Zoo, Jakarta, Indonesia

    [​IMG]
    @ZooGirlSD , San Diego Zoo, USA


    References

    Frantz et al. (2018): https://royalsocietypublishing.org/doi/10.1098/rspb.2017.2566

    Groves (1980): https://www.repository.naturalis.nl/document/150456

    Meijaard & Groves (2002): https://www.researchgate.net/public...f_babirusa_Babyrousa_sp_to_full_species_level
     
    Last edited: 21 Nov 2019
  14. Gondwana

    Gondwana Well-Known Member

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    There is a recent paper that incorporates a pretty solid data set of genetic and morphometric data for Babirusa: https://royalsocietypublishing.org/doi/10.1098/rspb.2017.2566

    The take home is that their genetic data support the presence of six population clusters, with the Moluccas (Buru) animals being most divergent (you have to look at the supplement rather than the main paper to see most of the genetic results). Their morphometric data show the Togian animals to be very different, with molar shape closer to some warty pigs than to other Babirusa.

    Good luck once you get to the Sus mess...
     
  15. lintworm

    lintworm Well-Known Member 15+ year member

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    Thanks, Sus is probably the most complicated genus of them all....

    Thanks for linking this study, I am surprised I missed such a big one, but given the title that is maybe not so strange. I will update the species profile to incorporate this information.

    It is a good study and although based on the morphological data, one could split togeanensis from babyrussa+celebensis, the genetic differences between the six identified group are quite small. The genetics do show that the 3 until now recognized taxa are monophyletic, but they are probably best treated as subspecies. There are also multiple potential subspecies on Sulawesi itself, but based on morphological data analysed in this study, differences are minor, even though they form identifiable genetic groups.
     
    Last edited: 21 Nov 2019
  16. UngulateNerd92

    UngulateNerd92 Well-Known Member Premium Member 5+ year member

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    Sorry to bump this thread, but I would like to notify you that the image you used intending to depict a female nominate Mule deer from ThylacineAlive at the Arizona Sonora Desert Museum is actually of a Coue's white tailed deer (Odocoileus virginianus couesi).
     
    lintworm, Gondwana and ThylacineAlive like this.
  17. lintworm

    lintworm Well-Known Member 15+ year member

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    Forest hog

    The Forest hog or Giant forest hog (Hylocherus meinertzhageni) is one of the very few discoveries by Richard Meinertzhagen that has not been debunked since. This is a large species of pig that occurs throughout much of tropical Africa from Liberia in the west to Kenya and Ethiopia in the east. Traditionally three subspecies have been recognized:

    H.m. meinertzhageni Mountains of the Albertine Rift in East DR Congo, Uganda, Rwanda, Uganda, Imatong Mountains in South Sudan, Kenya Highlands and presumably also in S Ethiopia
    H.m. rimator (includes ituriensis) Cameroon, Gabon, both Congo's and Central African Republic
    H.m. ivoriensis rainforests from Liberia to Ghana

    G&G elevate these three subspecies to full species.

    Sample sizes

    No sample sizes on skins are reported and no skin characteristics are given.

    Skulls (males / females)
    meinertzhageni 15-18/13-17
    rimator 7-10/5-6
    ituriensis 24-28/19-20
    ivoriensis 13-19/4-7

    Overall sample size is quite good, especially for males.

    Skins

    No skin characteristics given.

    Skulls

    Though there are some differences on average in the female sample, differences between the male samples are generally more pronounced. It is clear that meinertzhageni is larger than ivoriensis, with no to minimal overlap in several of the measurements: greatest skull length, palatal lengths, pre-orbital length and canine breadth. In the other measurements this difference is also easily visible. ituriensis + rimator fall in between of these measurements. On average meinertzhageni is still clearly larger and in some cases there is no overlap in measurements with rimator, but ituriensis is in between these two groups, but closer to rimator. In most measurements there is quite some overlap between ivoriensis and rimator and apparently they only separate in a multivariate analysis (of which results are not presented). Given the fact that there are very probable high correlations between the different measurements, it is questionable whether the results of a multivariate analysis are valid, as the measurements are not independent of each other.

    Additional data

    No genetic work on Forest hogs seems to have been done.

    Summarizing

    Based on multivariate analysis G&G split Forest hog in three species and although there are discrete differences between the western ivoriensis and the eastern meinertzhageni, the samples of Central Africa (rimator + ituriensis) fall in between. When looking at all measurements there seems to be a clear cline in size from ivoriensis to rimator to ituriensis to meinertzhageni stretching from west to east. It seems very premature to recognize multiple species in such a cline, especially when differences from one popullation to the next are relatively small, with the exception of meinertzhageni. The reason why meinertzhageni is bigger, could be because this species lives in cooler montane forests than the other species, which are mainly from lowland rainforests. Genetic research would be welcome, but current evidence seems to point more to a cline than to discrete species.

    Hylochoerus meinertzhageni meinertzhageni
    [​IMG]
    @Nick@Amsterdam , Uganda Wildlife Education Centre, Entebbe, Uganda

    Hylochoerus meinertzhageni rimator
    [​IMG]

    uploaded by @Joker1706 , Zoo Antwerpen, Belgium
     
  18. lintworm

    lintworm Well-Known Member 15+ year member

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    Philippine warty pig

    The Philippine warty pig (Sus philippensis s.l.) is a potential species complex of pigs in the Philippines. Traditionally one species was recognized with three subspecies:

    S.p. philippensis Luzon Island, Philippines
    S.p. mindanensis Mindanao Island, Philippines
    S.p. oliveri Mindoro Island, Philippines

    oliveri has been split by Groves in 2001 and this has classification has since been used by the IUCN, HMW and other sources.


    Sample size

    Skins are not mentioned in G&G (2011), but in Groves (1997) only 1 head of oliveri was described.

    Skulls
    philippensis 13-15
    mindanensis 6-9
    oliveri 3

    Teeth
    philippensis 5-8
    mindanensis 5-6
    oliveri 2-3

    Sample sizes are mostly very small for these taxa.


    Skins

    Groves (1997) description of the single oliveri head skin seems to give some minor differences with other taxa. There is however a lot of variation within the other taxa, so it is likely this variation also exists in oliveri and it is thus impossible to verify how distinct oliveri would be.


    Skulls

    oliveri is described as having a long palate and a wide bizygomatic breadth, compared to philippensis. When looking at the measurements there is however complete overlap with measurements of mindanensis and also wide overlap with philippensis. In multivariate analyses presented in Groves (1997) there is no difference between oliveri and the other two taxa.


    Teeth

    Several differences in length and breadth of different teeth are described for oliveri, but the measurements itself do largely overlap with the othr taxa. In multivariate analyses in Groves (1997) oliveri is not really distinct in maxillary dentition, but it does group apart when analysing mandibular dentition. It should however be noted that mindanensis and philippensis also group apart there, but with a smaller distance in between them.


    Additional data

    No studies on the genetic relationship of these taxa have been done as far as I am aware. Oliver (1995) mentions that mindanensis and philippensis have a karyotype of 2n=36, whereas oliveri has a karyotype of 2n=38.


    Summarizing

    Based on morphological data there is no ground to recognize oliveri as a distinct species, the described differences are either non-existent or very minor and all are based on extremely limited samples of oliveri. The difference in karyotype might however point to more variation than is currently recognized. For now I see no reason to prematurely split oliveri, but genetic research is of high priority here.

    [​IMG]
    @Nick@Amsterdam , Avilon Zoo, Philippines

    [​IMG]
    @Nick@Amsterdam , Avilon Zoo , Philippines

    References

    Groves (1997): Error - Cookies Turned Off

    Oliver (1995): http://mountainecology.org/index.php/me/article/viewFile/53/42
     
  19. lintworm

    lintworm Well-Known Member 15+ year member

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    Javan warty pig

    The Javan warty pig (Sus verrucosus) is a species of pig restricted to the islands of Java and Bawean in Indonesia. Tradtionally two subspecies were recognized:

    S.v. verrucosus Java, Indonesia
    S.v. blouchi Bawean, Indonesia

    G&G elevate blouchi to species status.


    Sample sizes

    No sample sizes are given by G&G, nor is any raw data presented, it might be that this data is presented in Groves (1981), to which I have no access. It is however likely to be a very low sample size


    Skins

    The colour of blouchi is described as yellowish, not reddish, which is the case for verrucosus


    Skulls

    Skulls of blouchi are described as much smaller than verrucosus with a skull length of 354 mm, whereas skull length in verrucosus is 408-429 mm.


    Additional data

    I have not been able to find any studies comparing dna of blouchi with verrucosus


    Summarizing

    Based on the extremely limited data available it seems very premature to elevate blouchi to species status. It might just be a small island morph of verrucosus, which seems just as likely an explanation to me as splitting it

    Sus verrucosus verrucosus
    [​IMG]
    @Vision , Gembira Loka Zoo, Yogyakarta, Indonesia

    [​IMG]
    @Vision , Gembira Loka Zoo, Yogyakarta, Indonesia

    Reference

    Groves (1981): CiNii 論文 - Ancestors for the pigs: Taxonomy and phylogeny of the genus Sus.
     
  20. lintworm

    lintworm Well-Known Member 15+ year member

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    Europe
    Eurasian wild pig


    The Eurasian wild pig or Wild boar (Sus scrofa) is one of the most widespread ungulate species, occuring throughout most of Europe and Asia, as well as a small population in North Africa. and occurs in a wide range of habitats. The Eurasian wild pig is a highly adaptable species and throughout its range there is a large morphological variation.The putative species Sus bucculentus has been shown to actually be an Eurasian wild pig. Traditionally around 19 subspecies have been recognized, which I have grouped here based on the groupings identified by Groves & Grubb (1993):

    Western
    S.s. scrofa W, SW & N Europa
    S.s. attila E Europe
    S.s. algirus Tunisia, Algeria and Morocco
    S.s. baeticus S Iberian Peninsula
    S.s. meridionalis Corsica & Sardinia (possibly introduced, subspecies potentially invalid)
    S.s. lybicus Bulgaria, Greece, Turkey & Levant
    S.s. majori C & S Italy
    S.s. nigripes Tianshan Mountains in Kyrgyzstan, NW China, Tajikistan and possibly N Iran

    Indian
    S.s. cristatus Himalaya S to Central India and east to Indochina
    S.s. affinis S India & Sri Lanka
    S.s. davidi E Iran to W India

    East Asian
    S.s. coreanus Korean peninsula
    S.s. leucomystax Japan
    S.s. moupinensis
    S.s. riukiuanus
    Ryukyu Islands (Japan)
    S.s. sibiricus Mongolia and E Russia
    S.s. taivanus Taiwan

    Southeast Asian
    S.s. vittatus Malay Peninsula, S of isthmus of Kra, and Indonesia archipelago

    G&G propose to recognize 13 species:
    S. scrofa (including attila, lybicus, algira, baeticus, majori)
    S. meridionalis
    S. nigripes
    S. sibiricus
    S. leucomystax
    S. taevanus
    S. riukiuanus
    S. davidi
    S. cristatus
    (includes affinis)
    S. vittatus
    S. moupinensis
    (for animalsof Myanmar & China, includes bucculentus)
    S. ussuricus (for animals of Heilongjiang, China, and the Far East)
    S. chirodontus (for animals of SC China)

    G&G state that splitting of meridionalis, nigripes (based on a single specimen), sibiricus and riukiuanus is preliminary, as their uniqueness could not be determined.

    Sample sizes

    For most splits G&G do not provide any evidence in their Ungulate Taxonomy book and they refer to a 1981 publication to which I do not have any access.

    They do however provide the skull data on which they base the split of moupinensis/chirodontus/ussuricus

    Skulls (males/females)
    moupinensis 36-41/4-6
    chirodontus 4-10/3
    ussuricus 3-8/0-5

    Skins

    No details on skins are given.

    Skulls

    Though G&G claim that their multivariate analysis shows a clear distinction between the taxa, the data presented in the book does not indicate any difference between moupinensis and chirodontus. ussuricus is distinct from the other taxa, as it is larger.

    taevanus and riukiuanus are both very small and G&G mention both taxa need to be compared closely with each other

    Additional data

    The Eurasian wild pig has been of great interest to researchers, not in the least because it is one of the most important domestic animals we have. There is plenty literature on the genetics of Eurasian wild pig, but data which can give a clear answer to whether the splits of G&G are valid are more scarce.

    The main paper looking at genetic differences between wild pigs across its range comes from Larson et al. (2005). They describe (Figure 1) a SE Asian origin of the Eurasian wild pig, something which has been backed up by multiple other studies. They show that vittatus is the most distinct taxon of them all. Additionally they find eight other genetic groups: Indochina, two in India, China + Korea + Japan + parts of Russia and the Stans, one in S China (might correspond to chirodontus?) Taiwan, Ryukyu Islands and finally Europe, W Asia& N Africa. The latter group consists of three smaller subgroups: Italy + Sardinia, W Asia and the rest of Europe + N Africa. Of all these groups one Indian group is most divergent

    Based on whole genomes Frantz et al. (2013) estimate the divergence between the European wild pig and the Asian wild pig 1 mya. The split between northern and southern Wild pigs they estimate at about 0.6 mya. There is however evidence of gene flow between these populations well after these splits. A study by Khalilzadeh et al. (2016) found that in Iran the European and Asian groups mix.

    Differences in karyotype have also been described, with 2n=36 being common in Europe and 2n=38 being the norm in Asia. Individuals with 2n=37 do however exist as well, so this might not be evidence of true speciation yet.

    Khederzadeh et al. (2018) analyzed the genetic differences between Wild pigs in Europe, W Asia and N Africa. They found that animals from the Caucasus were most distinct as were animals from the Italian peninsula. W European animals grouped together with the N African sample and these combined are somewhat distinct from the Eastern European & Balkan sample. These genetic results do not warrant the recognition of separate subspecies on the Iberian peninsula, this would mean castillianus and baeticus are synonyms of scrofa.

    Apollonio et al. (1988) describe that meridionalis of Sardinia is much smaller than pigs of the Italian mainland, but they cannot offer an explanation: both introgression with domestic pigs and island dwarfism are options. Genetically meriddionalis pigs are grouped with the Italian pigs (Larson et al. 2005).

    Summarizing

    There remains much to be learned about how many species of Eurasian wild pig exist. The only thing that is clear is that it is more than 1, but most likely less than described by G&G. There is quite good evidence to recognize recognize separate wild pig species at least in Europe + North Africa (scrofa, including attila, majori, lybicus and possibly meridionalis as subspecies and algirus as a synonym of scrofa), India (cristatus), SE Asia (vittatus) and at least 1, probably at least 2 and potentially at least four more species in Central and Eastern Asia. It is still unclear whether taevanus, riukiuanus and the different Chinese taxa should be treated. Based on dna results sibiricus, nigripes, davidi and leucomystax do not find any support for now, but extensive sampling in Asia is necessary. Then there is also the question on how the second Indian haplotype (could that be davidi) and the Indochina sample should be treated. More research is certainly needed to establish the phylogenetic relations in this species group and give a more informed answer on how many Eurasian wild pig species should be recognized. Currently I think we will eventually end up with 5-9 species of Eurasian wild pig, but surprises along the way are well possible.

    Sus (scrofa) scrofa scrofa
    [​IMG]

    @Daniel Sörensen , Wildpark Hannover, Germany

    [​IMG]
    @Maguari , Dierenrijk, Mierlo, Netherlands

    Sus (scrofa) scrofa lybicus
    [​IMG]

    @alexkant , Meir Segals Garden University Zoo, Israel

    Sus (scrofa) scrofa attilus
    [​IMG]
    @alexkant , Mogilev Zoo, Belarus

    Sus (scrofa) cristatus
    [​IMG]

    @Chlidonias , Wilpattu NP, Sri Lanka

    Sus (scrofa) vittatus
    [​IMG]

    @Chlidonias , Ujung Kulon NP, Indonesia

    [​IMG]
    @Vision , Pulau Ubin, Singapore

    Sus (scrofa) leucomystax leucomystax
    [​IMG]

    @Giant Eland , Tama Zoo, Japan

    Sus (scrofa) leucomystax moupinensis
    [​IMG]

    @Deer Forest , Tangjiahe NNR, China

    Animal from Myanmar
    [​IMG]
    @Chlidonias , Yadanabon Zoo, Myanmar

    Animal from Iran
    [​IMG]
    @fofo , Tehran Zoo, Iran

    Animals from Vietnam
    [​IMG]
    @Maguari , Saigon Zoo, Vietnam


    References

    Apollonio et al. (1988): https://www.tandfonline.com/doi/pdf/10.1080/11250008809386619

    Frantz et al. (2013): Genome sequencing reveals fine scale diversification and reticulation history during speciation in Sus

    Groves & Grubb (1993): http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.693.1866&rep=rep1&type=pdf#page=120

    Khalilzadeh et al. (2016): Contact Zone of Asian and European Wild Boar at North West of Iran

    Khederzadeh et al. (2018): Error - Cookies Turned Off

    Larson et al. (2005): Worldwide Phylogeography of Wild Boar Reveals Multiple Centers of Pig Domestication