Pretty sure the forum's already proven that Fossa are just Thylacines in disguise (or was it the other way around
) but I'd be happy to accept a research grant to do see for myself 
~Thylo
Ungulate taxonomy revisited: the evidence for the splits of G&G
- Thread starter lintworm
- Start date
Pretty sure the forum's already proven that Fossa are just Thylacines in disguise (or was it the other way around
) but I'd be happy to accept a research grant to do see for myself ~Thylo
Saiga
The Saiga (Saiga tartarica) is a unique antelope from the steppes of Eurasia. Especially during the Ice Ages it was very widespread and even in historical times it occured from the Carpathian mountains to Mongolia. Currently its distribution is limited to remnants in Mongolia, Kazakhstan, Russia, Uzbekistan and Turkmenistan. Traditionally two subspecies have been recognized:
S.t. tartarica Kazakhstan, Russia, Uzbekistan & Turkmenistan
S.t. mongolica Mongolia
G&G elevate both subspecies to species status
Sample sizes
No sample sizes are given and G&G say that the characters of the two very distinctive taxa largely follow Bannikov (1963). Whether this means all the raw data is taken from Bannikov or whether the data from G&G gives the same results as Bannikov is unclear. I do however not have access to Bannikov (1963), so I cannot tell...
Skins
The tartarica summer coat is described as yellowish-red, paler on the flanks and with white undersides. Darker zones are present on the shoulders and the loins. The mongolica summer coat is described as sandy gray, with a dorsal region that is not darkened, but with a brown spot on the lumbar region that is large and sharply bordered. The difference in pelage color yellowish-red vs. sandy gray is however completely invisible in the pictures provided in Castello's field guide, though the belly of mongolica is less whitish. The winter coat of tartarica is described as very light gray coloured, the winter coat of mongolica is not described.
Skulls & horns
Skull length is 222-250 mm in tartarica males, 205-209 in tartarica females and 203-237 in mongolica (unspecified whether male or female). There is thus no clear difference in skull length. Horns of tartarica are longer (minimum length 280 mm) than mongolica (maximum length 220 mm). Thickness of the horns seems quite similar 25-33 mm in tartarica vs. up to 28 mm in mongolica, though G&G state that mongolica has thinner horns (but that is probably just based on averages). The rings on the horns are more strongly marked in tartarica than in mongolica. The nasal opening is described as more raised in tartarica than in mongolica.
Additional data
Kholodova et al. (2006) find a slight but clear genetic differentiation between the two taxa. The differences are however much smaller than in African buffalo, Grant's gazelle or Kob. The estimated time of divergence between the two populations is likely to have been in the late pleistocene or even the early holocene, so is very recent.
Summarizing
Though there are some differences in the horns, the rest of the described differences are rather minor and I do not know the sample sizes that were involved. Combined with the genetic data that indicate rather small differences and a very recent date of divergence, I do not see good reasons to elevate the subspecies to species status. If you would be a strict follower of the PSC concept both the morphological and genetic data would allow a split, but strictly following only 1 species concept never has been the smartest idea, especially as several populations of tartarica should then also be elevated to species status.
Saiga tartarica tartarica
Male in winter coat
@Zebraduiker, Cologne Zoo, Germany
Male in summer coat
@Parrotsandrew , Edinburgh Zoo, UK
Female in summer coat:
@Arizona Docent , San Diego Zoo, USA
no pictures of mongolica have been uploaded to the gallery yet.
References
Bannikov, A.G. 1963. Die Saiga-Antilope (Saiga tartarica L.). Die Neue Brehm-Buecherei No. 320. Wittenberg-Lutherstadt, Germany: A. Ziemsen.
Khodolova et al. (2006): https://www.iccs.org.uk/wp-content/papers/Kholodova2000.pdf
Next: Goitered gazelle
The Saiga (Saiga tartarica) is a unique antelope from the steppes of Eurasia. Especially during the Ice Ages it was very widespread and even in historical times it occured from the Carpathian mountains to Mongolia. Currently its distribution is limited to remnants in Mongolia, Kazakhstan, Russia, Uzbekistan and Turkmenistan. Traditionally two subspecies have been recognized:
S.t. tartarica Kazakhstan, Russia, Uzbekistan & Turkmenistan
S.t. mongolica Mongolia
G&G elevate both subspecies to species status
Sample sizes
No sample sizes are given and G&G say that the characters of the two very distinctive taxa largely follow Bannikov (1963). Whether this means all the raw data is taken from Bannikov or whether the data from G&G gives the same results as Bannikov is unclear. I do however not have access to Bannikov (1963), so I cannot tell...
Skins
The tartarica summer coat is described as yellowish-red, paler on the flanks and with white undersides. Darker zones are present on the shoulders and the loins. The mongolica summer coat is described as sandy gray, with a dorsal region that is not darkened, but with a brown spot on the lumbar region that is large and sharply bordered. The difference in pelage color yellowish-red vs. sandy gray is however completely invisible in the pictures provided in Castello's field guide, though the belly of mongolica is less whitish. The winter coat of tartarica is described as very light gray coloured, the winter coat of mongolica is not described.
Skulls & horns
Skull length is 222-250 mm in tartarica males, 205-209 in tartarica females and 203-237 in mongolica (unspecified whether male or female). There is thus no clear difference in skull length. Horns of tartarica are longer (minimum length 280 mm) than mongolica (maximum length 220 mm). Thickness of the horns seems quite similar 25-33 mm in tartarica vs. up to 28 mm in mongolica, though G&G state that mongolica has thinner horns (but that is probably just based on averages). The rings on the horns are more strongly marked in tartarica than in mongolica. The nasal opening is described as more raised in tartarica than in mongolica.
Additional data
Kholodova et al. (2006) find a slight but clear genetic differentiation between the two taxa. The differences are however much smaller than in African buffalo, Grant's gazelle or Kob. The estimated time of divergence between the two populations is likely to have been in the late pleistocene or even the early holocene, so is very recent.
Summarizing
Though there are some differences in the horns, the rest of the described differences are rather minor and I do not know the sample sizes that were involved. Combined with the genetic data that indicate rather small differences and a very recent date of divergence, I do not see good reasons to elevate the subspecies to species status. If you would be a strict follower of the PSC concept both the morphological and genetic data would allow a split, but strictly following only 1 species concept never has been the smartest idea, especially as several populations of tartarica should then also be elevated to species status.
Saiga tartarica tartarica
Male in winter coat
@Zebraduiker, Cologne Zoo, Germany
Male in summer coat
@Parrotsandrew , Edinburgh Zoo, UK
Female in summer coat:
@Arizona Docent , San Diego Zoo, USA
no pictures of mongolica have been uploaded to the gallery yet.
References
Bannikov, A.G. 1963. Die Saiga-Antilope (Saiga tartarica L.). Die Neue Brehm-Buecherei No. 320. Wittenberg-Lutherstadt, Germany: A. Ziemsen.
Khodolova et al. (2006): https://www.iccs.org.uk/wp-content/papers/Kholodova2000.pdf
Next: Goitered gazelle
Last edited:
Goitered gazelle
The Goitered gazelle (Gazella subgutturosa) (sometimes called Goitred gazelle) is a small gazelle that occurs from Azerbaijan through Iraq and Iran to NW China and Mongolia. The Sand gazelle (Gazella marica) was formerly classified as a subspecies of the Goitered gazelle and is treated by G&G as sister species of the Goitered gazelle. The Sand gazelle is however most closely related to the Slender-horned gazelle (Gazella leptoceros) and the Cuvier's gazelle (Gazella cuvieri) based on genetic data (Wacher et al. 2010; Hassanin et al. 2012). Interestingly female Goitered gazelles are generally hornless, whereas Sand gazelle females are not and their horns closely resemble those of Slender-horned gazelles. Apart from the Sand gazelle three subspecies were generally recognized:
G.s. subgutturosa from Azerbaijan, S Turkey and Iraq to Iran and Kazakhstan, Turkmenistan, Uzbekistan
G.s. hillieriana Mongolia and N and C China
G.s. yarkandensis extreme W China
G&G recognize four species: hillieriana is lumped within yarkandensis and yarkandensis is elevated to species status. subgutturosa is split into 3 species: subgutturosa from Iran E of the Zagros mountains and possibly Azerbaijan, gracilicornis from Kazakhstan, Turkmenistan and Uzbekistan and an undescribed species from Iraq, Anatolia and Iran W of the Zagros mountains.
Sample sizes
No sample sizes for skins are given.
The table in which horn and skull characteristics are summarized is a bit of a mess. Though it is clear to which taxa hillieriana, yarkandensis, "subgutturosa Azerbaijan" and Iraq belong. Iran seistanica must refer to subgutturosa and diversicornis is probably an error and should be gracilicornis. diversicornis is a subspecies of the Przewalski's gazelle (Procapra przewalskii).
Skulls (Males only)
Iraq 2-6
subgutturosa:
Caucasus 2
Iran 6-8
gracilicornis 12-14
yarkandensis 8-10
hillieriana 19-25
Horns (Males only)
Iraq 7-10
subgutturosa:
Caucasus 1
Iran 9-10
gracilicornis 17
yarkandensis 7-9
hillieriana 18-25
Sample sizes are relatively limited for most taxa, though not extremely low when combining regions.
Skins
yarkandensis and gracilicornis are very similar in color being sandy-brown to sandy-grey in summer and lighter in winter. subgutturosa is described as reddish or grayish-sandy above. All taxa have white below and on the buttocks. All three taxa have an indistinct flank stripe and facial stripes, which are apparently more strongly marked in the Iraq specimens.
Horns & Skulls
Though on average there are some minimal differences between the taxa in skull measurements, yarkandensis, gracilicornis and subgutturosa measurements overlap widely for all four measurements given. The only thing is that the 2 Caucasus samples are somewhat larger than most averages, though fall within measurements of the other taxa. The Iraqi samples do however have narrower posterial nasal measurements, though there is some overlap. The two measurements for Gt 1 were also smaller than the other taxa, but sample size is minimal. In horn measurements the averages between the taxa differ, there is however wide overlap between the different taxa in all measurements. The only difference is that the basal breadth is distinctly larger in gracilicornis when compared to subgutturosa and the Iraqi samples. yarkandensis does however overlap completely with subgutturosa, gracilicornis and the Iraqi samples. Tip to tip distance is on average much smaller in yarkandensis when compared to gracilicornis and the Irani samples of subgutturosa, but there is wide overlap in the tip-to-tip distance between these taxa and the subgutturosa from the Caucasus are the same as yarkandensis in tip to tip distance.
Additional data
Hayatgheib et al. (2011) conclude that the gazelles from Iran W of the Zagros mountains and Iraq are very similar and smaller than the gazelles from E of the Zagros mountains. There models are however extremely overfitted given the sample sizes are extremely small. Looking at the table with the individual measurements there does however seem to be a trend that the Iraqi and W Iranian samples are smaller than the samples of E of the Zagros mountains. But with the very small sample sizes (12 samples were included in the whole paper and there was no distinguishing between males and females) no conclusions can be drawn. Apart from that Heptner et al. (1961) describe that the Caucasus animals are darker coloured with a clear dark flank stripe and heavy tail tuft.
Summarizing
Though G&G make some hard statements of gracilicornis being very wide acros the base of the horns or that yarkandensis has a long skull or that the Iraqi samples are much smaller, the provided evidence is not convincing. What is interesting though is that both their data and the data from Hayatgheib suggest that there might be an undescribed Goitered gazelle taxa W of the Zagros mountains. The animals from the Caucasus might also warrant subspecies status but given the almost complete lack of data from this region, no conclusions can be reached. There does however seem to be nothing wrong with lumping yarkandensis and hillieriana. Based on the skins the Goitered gazelles from Kazakhstan, Turkmenistan and Uzbekistan might better be treated as yarkandensis than as subgutturosa. For now that would mean that there are only 2 described subspecies: subgutturosa and yarkandensis + an undescribed subspecies from W Iran and Iraq.
Gazella subgutturosa subgutturosa
Male:
@Giant Eland , Miami Zoo, USA
Female:
@Morgan, ZooParc Overloon, Netherlands
Gazella subgutturosa yarkandensis
@Deer Forest , Beijing Zoo, China
Goitered gazelle from Antalya Zoo, Turkey, could be of the undescribed subspecies, if sourced in the country:
@Zaz , Antalya Zoo, Turkey
References:
Hassanin et al. (2012): https://www.sciencedirect.com/science/article/pii/S1631069111002800
Hayatgheib et al. (2011): https://ijer.ut.ac.ir/article_307_e1d392ccb2de9eb2f4438f1395ec990e.pdf
Heptner, V., A.A, Nasimovich, A.G. Bannikov. 1961. Artiodactyla and Perrisodactyla. Vol. 1 Mammals of the Soviet Union. Moscow: Vysshava Shkola.
Wacher et al. (2010): https://www.researchgate.net/profil...bgutturosa/links/09e41508a9e33eb89f000000.pdf
Next: Arabian gazelle
The Goitered gazelle (Gazella subgutturosa) (sometimes called Goitred gazelle) is a small gazelle that occurs from Azerbaijan through Iraq and Iran to NW China and Mongolia. The Sand gazelle (Gazella marica) was formerly classified as a subspecies of the Goitered gazelle and is treated by G&G as sister species of the Goitered gazelle. The Sand gazelle is however most closely related to the Slender-horned gazelle (Gazella leptoceros) and the Cuvier's gazelle (Gazella cuvieri) based on genetic data (Wacher et al. 2010; Hassanin et al. 2012). Interestingly female Goitered gazelles are generally hornless, whereas Sand gazelle females are not and their horns closely resemble those of Slender-horned gazelles. Apart from the Sand gazelle three subspecies were generally recognized:
G.s. subgutturosa from Azerbaijan, S Turkey and Iraq to Iran and Kazakhstan, Turkmenistan, Uzbekistan
G.s. hillieriana Mongolia and N and C China
G.s. yarkandensis extreme W China
G&G recognize four species: hillieriana is lumped within yarkandensis and yarkandensis is elevated to species status. subgutturosa is split into 3 species: subgutturosa from Iran E of the Zagros mountains and possibly Azerbaijan, gracilicornis from Kazakhstan, Turkmenistan and Uzbekistan and an undescribed species from Iraq, Anatolia and Iran W of the Zagros mountains.
Sample sizes
No sample sizes for skins are given.
The table in which horn and skull characteristics are summarized is a bit of a mess. Though it is clear to which taxa hillieriana, yarkandensis, "subgutturosa Azerbaijan" and Iraq belong. Iran seistanica must refer to subgutturosa and diversicornis is probably an error and should be gracilicornis. diversicornis is a subspecies of the Przewalski's gazelle (Procapra przewalskii).
Skulls (Males only)
Iraq 2-6
subgutturosa:
Caucasus 2
Iran 6-8
gracilicornis 12-14
yarkandensis 8-10
hillieriana 19-25
Horns (Males only)
Iraq 7-10
subgutturosa:
Caucasus 1
Iran 9-10
gracilicornis 17
yarkandensis 7-9
hillieriana 18-25
Sample sizes are relatively limited for most taxa, though not extremely low when combining regions.
Skins
yarkandensis and gracilicornis are very similar in color being sandy-brown to sandy-grey in summer and lighter in winter. subgutturosa is described as reddish or grayish-sandy above. All taxa have white below and on the buttocks. All three taxa have an indistinct flank stripe and facial stripes, which are apparently more strongly marked in the Iraq specimens.
Horns & Skulls
Though on average there are some minimal differences between the taxa in skull measurements, yarkandensis, gracilicornis and subgutturosa measurements overlap widely for all four measurements given. The only thing is that the 2 Caucasus samples are somewhat larger than most averages, though fall within measurements of the other taxa. The Iraqi samples do however have narrower posterial nasal measurements, though there is some overlap. The two measurements for Gt 1 were also smaller than the other taxa, but sample size is minimal. In horn measurements the averages between the taxa differ, there is however wide overlap between the different taxa in all measurements. The only difference is that the basal breadth is distinctly larger in gracilicornis when compared to subgutturosa and the Iraqi samples. yarkandensis does however overlap completely with subgutturosa, gracilicornis and the Iraqi samples. Tip to tip distance is on average much smaller in yarkandensis when compared to gracilicornis and the Irani samples of subgutturosa, but there is wide overlap in the tip-to-tip distance between these taxa and the subgutturosa from the Caucasus are the same as yarkandensis in tip to tip distance.
Additional data
Hayatgheib et al. (2011) conclude that the gazelles from Iran W of the Zagros mountains and Iraq are very similar and smaller than the gazelles from E of the Zagros mountains. There models are however extremely overfitted given the sample sizes are extremely small. Looking at the table with the individual measurements there does however seem to be a trend that the Iraqi and W Iranian samples are smaller than the samples of E of the Zagros mountains. But with the very small sample sizes (12 samples were included in the whole paper and there was no distinguishing between males and females) no conclusions can be drawn. Apart from that Heptner et al. (1961) describe that the Caucasus animals are darker coloured with a clear dark flank stripe and heavy tail tuft.
Summarizing
Though G&G make some hard statements of gracilicornis being very wide acros the base of the horns or that yarkandensis has a long skull or that the Iraqi samples are much smaller, the provided evidence is not convincing. What is interesting though is that both their data and the data from Hayatgheib suggest that there might be an undescribed Goitered gazelle taxa W of the Zagros mountains. The animals from the Caucasus might also warrant subspecies status but given the almost complete lack of data from this region, no conclusions can be reached. There does however seem to be nothing wrong with lumping yarkandensis and hillieriana. Based on the skins the Goitered gazelles from Kazakhstan, Turkmenistan and Uzbekistan might better be treated as yarkandensis than as subgutturosa. For now that would mean that there are only 2 described subspecies: subgutturosa and yarkandensis + an undescribed subspecies from W Iran and Iraq.
Gazella subgutturosa subgutturosa
Male:
@Giant Eland , Miami Zoo, USA
Female:
@Morgan, ZooParc Overloon, Netherlands
Gazella subgutturosa yarkandensis
@Deer Forest , Beijing Zoo, China
Goitered gazelle from Antalya Zoo, Turkey, could be of the undescribed subspecies, if sourced in the country:
@Zaz , Antalya Zoo, Turkey
References:
Hassanin et al. (2012): https://www.sciencedirect.com/science/article/pii/S1631069111002800
Hayatgheib et al. (2011): https://ijer.ut.ac.ir/article_307_e1d392ccb2de9eb2f4438f1395ec990e.pdf
Heptner, V., A.A, Nasimovich, A.G. Bannikov. 1961. Artiodactyla and Perrisodactyla. Vol. 1 Mammals of the Soviet Union. Moscow: Vysshava Shkola.
Wacher et al. (2010): https://www.researchgate.net/profil...bgutturosa/links/09e41508a9e33eb89f000000.pdf
Next: Arabian gazelle
Last edited:
Arabian gazelle
The Arabian gazelle (Gazella arabica) was formerly lumped with the Mountain gazelle (Gazella gazella). Genetic data did however indicate a clear split between Gazella gazella from C & N Israel and the Golan heights and the remaining Gazella from the Arabian peninsula (Baermann et al. 2012; Lerp et al. 2013). The remaining subspecies that were formerly lumped with gazella are now treated as Gazella arabica by the IUCN. There has always been confusion about which gazelles belong to which species on the Arabian peninsula and there has been confusion with G. marica, G. dorcas and the now extinct G. saudiya. To add to the confusion Baermann et al. (2012) found that the type specimen of arabica, a skull and two skins belong to two different species. The skull was said to come from the Farasan Islands SW of Arabia, but it was already noted that it was quite dissimilar from the gazelles currently found there. The skins were apparently collected from the Sinai. Genetic data indicated that the skull was nested well within the "northern" Gazella gazella and could thus impossibly be from Farasan Island, but was from Israel. The skins nested well within the "southern" arabica clade. It is now debatable whether the arabica should be treated as a junior synonym to gazella and that the real arabica should be called cora or whether the southern species can continued to be called arabica with cora as a synonym. As IUCN lists the gazelles as Gazella arabica I will do so too. The whole story can be read (open access) in Baermann et al. (2012).
Then on to the subspecies. Wilson & Reeder recognize 5 subspecies in the Arabian gazelle:
G.a. cora = arabica isolated populations throughout the Arabian peninsula in Oman, Yemen, Saudi Arabia, United Arab Emirates.
G.a. acaciae Aravi Rift Valley, Israel
G.a. dareshurii Farrur Island
G.a. farasani Great Farasan and Zifaf Island
G.a. muscatensis NW Oman
G&G recognize 7 species (excluding Gazella arabica which they base purely on the skull which has now been shown to be a Gazella gazella). cora, acaciae, dareshurii, muscatensis are all elevated to species level. Populations of cora of SW Saudi Arabia and SW Yemen are elevated to species status and called erlangeri with farasani as subspecies and animals from Great Hanish Island as an additional subspecies: hanishi. Additionally karamii is described from Bushehr, Iran and bilkis as a possibly extinct taxa from Yemen. in HMW bilkis is not recognized.
Sample sizes
Exact sample sizes for skins are not given, but are extremely minimal for karamii, dareshurii and bilkis.
Horns & skulls (males only)
acaciae 3-5
cora 21-27
dareshurii 7-11
erlangeri 5
farasani 3-12
karamii 1
muscatensis 1-6
hanishi 1-2 (based on literature description)
bilkis Not clearly summarized, but very possible 2
Annoyingly bilkis is not included in the overview table of measurements, neither is hanishi. Additionally for muscatensis no range is given for all measurements nor a standard deviation.
Except for dareshurii and cora sample sizes are very small.
Skins
No skins were seen for dareshurii only photographs which show a pale sandy brown animal. karamii, erlangeri and muscatensis are very similar with a very dark ground colour, just as bilkis. cora is more lightly coloured, though darker individuals also occur (the UK animals for example). acaciae is described as dark earth brown. Additionally there are some minor differences with regards to the nose spot, facial colour and the flank and pygal stripes.
Skulls & horns
G&G claim that the three desert taxa acaciae, cora and erlangeri are distinct on multivariate analysis. They did however completely overfit their data as there are 1.5-2x as many characteristics scored than samples available for both acaciae and erlangeri. Looking at the skull measurements between the 3 taxa there is very wide overlap in all characteristics. Measurements of dareshurii also fall widely within the range of these taxa, as do muscatensis, karamii and farasani. No skull measurements at all are available for hanishi and bilkis
In terms of horns the 2 bilkis samples fall almost completely within the variation of cora and erlangeri, though one of the tip-to-tip distance measures is slightly smaller than any of the other individuals measured. hanishi also falls within the variation of the other taxa. In basically all measurements given in their table there is wide overlap between the taxa and no conclusions can be drawn whatsoever based on most measurements. The only things are that muscatensis horns are shorter than acaciae, but as no range is given for muscatensis it is unclear whether there is overlap in measurements. farasani has a very small horn span, which is smaller than any of the other taxa with probably minimal overlap (but again, no range of measurements given). The one karamii sample has an extremely large tip-to-tip distance and horn span, much larger than any other sample. This animal did however live in the Berlin Zoo in the 1920s and it is common for zoo gazelles to have deformed horns (see for example Slender-horned gazelle), so I don't think those measurements can be seen as representative for the region.
Additional data
The genetic data used for the split between G. arabica and G. gazella did only show very minor genetic differences between all the G. arabica samples. Even though many samples came from zoos or breeding centers, but based on them there is no reason whatsoever to suspect hidden variation in G. arabica. Wronski et al. 2017 found that erlangeri was nested well within arabica.
Summarizing
This is G&G at it's worst with multiple species described based on extremely limited samples, sometimes even only 1 skull + skin. I therefore see no reason to elevate any of these taxa to species status and there are probably too many subspecies recognized anyway. Even though there are some small differences in pelage and in rare cases in the skulls. The taxa that are most distinct and would surely warrant subspecies status would be farasani, acaciae, the nominate cora/arabica and possibly muscatensis and dareshurii. I am not convinced at all that erlangeri is really distinct and the evidence for karamii, hanishi and bilkis as distinct taxa is extremely limited as well. It would be interesting to see whether scientists less busy with oversplitting would be able to find subspecies delimitations in this species.
Gazella arabica arabica (=G.a. cora)
@gentle lemur , Chester Zoo, UK
Gazella arabica acaciae
@bongowwf , Negev Desert, Israel
No pictures have been uploaded of the other taxa.
References
Baermann et al. (2012): https://www.sciencedirect.com/science/article/pii/S1616504712000845
Lerp et al. (2013): https://s3.amazonaws.com/academia.e...=Phylogenetic_and_population_genetic_anal.pdf
Wronski et al. (2017) Dark grey gazelles Gazella (Cetartiodactyla: Bovidae) in Arabia: Threatened species or domestic pet?
Next: Indian gazelle
The Arabian gazelle (Gazella arabica) was formerly lumped with the Mountain gazelle (Gazella gazella). Genetic data did however indicate a clear split between Gazella gazella from C & N Israel and the Golan heights and the remaining Gazella from the Arabian peninsula (Baermann et al. 2012; Lerp et al. 2013). The remaining subspecies that were formerly lumped with gazella are now treated as Gazella arabica by the IUCN. There has always been confusion about which gazelles belong to which species on the Arabian peninsula and there has been confusion with G. marica, G. dorcas and the now extinct G. saudiya. To add to the confusion Baermann et al. (2012) found that the type specimen of arabica, a skull and two skins belong to two different species. The skull was said to come from the Farasan Islands SW of Arabia, but it was already noted that it was quite dissimilar from the gazelles currently found there. The skins were apparently collected from the Sinai. Genetic data indicated that the skull was nested well within the "northern" Gazella gazella and could thus impossibly be from Farasan Island, but was from Israel. The skins nested well within the "southern" arabica clade. It is now debatable whether the arabica should be treated as a junior synonym to gazella and that the real arabica should be called cora or whether the southern species can continued to be called arabica with cora as a synonym. As IUCN lists the gazelles as Gazella arabica I will do so too. The whole story can be read (open access) in Baermann et al. (2012).
Then on to the subspecies. Wilson & Reeder recognize 5 subspecies in the Arabian gazelle:
G.a. cora = arabica isolated populations throughout the Arabian peninsula in Oman, Yemen, Saudi Arabia, United Arab Emirates.
G.a. acaciae Aravi Rift Valley, Israel
G.a. dareshurii Farrur Island
G.a. farasani Great Farasan and Zifaf Island
G.a. muscatensis NW Oman
G&G recognize 7 species (excluding Gazella arabica which they base purely on the skull which has now been shown to be a Gazella gazella). cora, acaciae, dareshurii, muscatensis are all elevated to species level. Populations of cora of SW Saudi Arabia and SW Yemen are elevated to species status and called erlangeri with farasani as subspecies and animals from Great Hanish Island as an additional subspecies: hanishi. Additionally karamii is described from Bushehr, Iran and bilkis as a possibly extinct taxa from Yemen. in HMW bilkis is not recognized.
Sample sizes
Exact sample sizes for skins are not given, but are extremely minimal for karamii, dareshurii and bilkis.
Horns & skulls (males only)
acaciae 3-5
cora 21-27
dareshurii 7-11
erlangeri 5
farasani 3-12
karamii 1
muscatensis 1-6
hanishi 1-2 (based on literature description)
bilkis Not clearly summarized, but very possible 2
Annoyingly bilkis is not included in the overview table of measurements, neither is hanishi. Additionally for muscatensis no range is given for all measurements nor a standard deviation.
Except for dareshurii and cora sample sizes are very small.
Skins
No skins were seen for dareshurii only photographs which show a pale sandy brown animal. karamii, erlangeri and muscatensis are very similar with a very dark ground colour, just as bilkis. cora is more lightly coloured, though darker individuals also occur (the UK animals for example). acaciae is described as dark earth brown. Additionally there are some minor differences with regards to the nose spot, facial colour and the flank and pygal stripes.
Skulls & horns
G&G claim that the three desert taxa acaciae, cora and erlangeri are distinct on multivariate analysis. They did however completely overfit their data as there are 1.5-2x as many characteristics scored than samples available for both acaciae and erlangeri. Looking at the skull measurements between the 3 taxa there is very wide overlap in all characteristics. Measurements of dareshurii also fall widely within the range of these taxa, as do muscatensis, karamii and farasani. No skull measurements at all are available for hanishi and bilkis
In terms of horns the 2 bilkis samples fall almost completely within the variation of cora and erlangeri, though one of the tip-to-tip distance measures is slightly smaller than any of the other individuals measured. hanishi also falls within the variation of the other taxa. In basically all measurements given in their table there is wide overlap between the taxa and no conclusions can be drawn whatsoever based on most measurements. The only things are that muscatensis horns are shorter than acaciae, but as no range is given for muscatensis it is unclear whether there is overlap in measurements. farasani has a very small horn span, which is smaller than any of the other taxa with probably minimal overlap (but again, no range of measurements given). The one karamii sample has an extremely large tip-to-tip distance and horn span, much larger than any other sample. This animal did however live in the Berlin Zoo in the 1920s and it is common for zoo gazelles to have deformed horns (see for example Slender-horned gazelle), so I don't think those measurements can be seen as representative for the region.
Additional data
The genetic data used for the split between G. arabica and G. gazella did only show very minor genetic differences between all the G. arabica samples. Even though many samples came from zoos or breeding centers, but based on them there is no reason whatsoever to suspect hidden variation in G. arabica. Wronski et al. 2017 found that erlangeri was nested well within arabica.
Summarizing
This is G&G at it's worst with multiple species described based on extremely limited samples, sometimes even only 1 skull + skin. I therefore see no reason to elevate any of these taxa to species status and there are probably too many subspecies recognized anyway. Even though there are some small differences in pelage and in rare cases in the skulls. The taxa that are most distinct and would surely warrant subspecies status would be farasani, acaciae, the nominate cora/arabica and possibly muscatensis and dareshurii. I am not convinced at all that erlangeri is really distinct and the evidence for karamii, hanishi and bilkis as distinct taxa is extremely limited as well. It would be interesting to see whether scientists less busy with oversplitting would be able to find subspecies delimitations in this species.
Gazella arabica arabica (=G.a. cora)
@gentle lemur , Chester Zoo, UK
Gazella arabica acaciae
@bongowwf , Negev Desert, Israel
No pictures have been uploaded of the other taxa.
References
Baermann et al. (2012): https://www.sciencedirect.com/science/article/pii/S1616504712000845
Lerp et al. (2013): https://s3.amazonaws.com/academia.e...=Phylogenetic_and_population_genetic_anal.pdf
Wronski et al. (2017) Dark grey gazelles Gazella (Cetartiodactyla: Bovidae) in Arabia: Threatened species or domestic pet?
Next: Indian gazelle
Yep that is pretty amazing indeed. A large part of the genetic research comes from 1 Danish project led by Arctander and Siegismund. There is so much work to do with regards to bovid genetics and hopefully G&G have stirred things up a little as a lot of their splits can probably easily be disproven with genetic work.
Indian gazelle
The Indian gazelle or Chinkara (Gazella bennettii) is an Asian species of Gazelle that occurs from Iran, through Pakistan to India. Currently 5 subspecies are generally recognized, though karami, treated here as a subspecies of arabica is sometimes also listed as a subspecies, though morphologically it is much more similar to arabica.
G.b. bennettii Decca Plateau & Ganges Valley
G.b. shikarii Iran from Tehran to Touran Biosphere Reserve
G.b. fuscifrons SE Iran and into the Makran coastal region of Pakistan
G.b. christyi Thar desert, Gujarat, Rajashtan
G.b. salinarum Salt range and E to Delhi
Naturally G&G elevate all subspecies to species status
Sample sizes
Sample sizes for skins are not given
Skulls/horns (Males/females)
bennettii 8-20/3-10
christyi 7-33/1-4
fuscifrons 15-22/6-11
salinarum 7-15/1-3
shikarii 3-4/1-2
Except for shikarii the other taxa have relatively ok sample sizes for males, but on the small side for multivariate analyses with 10 variables which was performed (exact results are not given). For females sample sizes range from extremely small to okish, but still too small for any multivariate analyses.
Skins
shikarii is reported as having a light skin colour, whereas fuscifrons is said to be dark, bennettii is said to be dull reddish brown, christyi very pale drab brown and salinarum a rich tobacco brown. salinarum also has no contrasting flank band, which is said to be prominent in fuscifrons, christyi has slightly darker median dorsal and lower flank zones, which are clearly more tawny and abrubtly darker in bennettii. No dorsal band presence/absence is described for shikarii, but based on pictures there is one present very slightly. A nose spot is described only for fuscifrons, though christyi, salinarum and bennettii also clearly have one based on pictures in Castello's field guide. Only shikarii has a less pronounced spot.
Skulls & horns
Female horns are described to be very long in shikarii, but there are only 2 datapoints which fall completely in the variation of fuscifrons. Horn divergence (which I assume corresponds with their horn span measurement) is said to be largest in shikarii, but though true on average the variation falls completely within the other taxa and 3 of the 4 other taxa have higher maximum spans in males, for females there are hardly any data points available here. fuscifrons is described as having a small skull, especially for females, but this is not reflected in the data at all. The horns are also said to be much less divergent, but the variation falls completely within the other taxa. bennettii females are said to have larger nasals, but though true on average, the overlap in measurements is huge with the other taxa. The reported less divergent horns in bennettii males compared to the span is also not reflected in the data. The reported differences for christyi are also not represented in the data. The same goes for the reported unique characteristics of salinarum, though the horns are indeed slightly less turned inside than the other taxa, except bennettii, but see picture of fuscifrons below and in Castello's field guide, which suggest that this characteristic is also there in fuscifrons.
Additional data
I am not aware of any research looking at the genetical differences between the reported subspecies.
Summarizing
Overall I see no reason to recognize any of the proposed splits. There are absolutely no clear differences in the skull and horn measurements and though there are some slight differences between the skin, especially in overall colour and how prominent the flank band is, that is far too little to warrant species status.
Gazella bennettii (likely bennettii)
@Giant Eland , New Delhi Zoo, India
Gazella bennettii (likely fuscifrons)
@J I N X , Peshawar Zoo, Pakistan
Gazella bennettii shikarii
@fofo , Iran
Gazella bennetti christyi
@toto98 , Thar Desert, India
No pictures of salinarum have been uploaded yet to the gallery.
next: the remaining Antilopini.
The Indian gazelle or Chinkara (Gazella bennettii) is an Asian species of Gazelle that occurs from Iran, through Pakistan to India. Currently 5 subspecies are generally recognized, though karami, treated here as a subspecies of arabica is sometimes also listed as a subspecies, though morphologically it is much more similar to arabica.
G.b. bennettii Decca Plateau & Ganges Valley
G.b. shikarii Iran from Tehran to Touran Biosphere Reserve
G.b. fuscifrons SE Iran and into the Makran coastal region of Pakistan
G.b. christyi Thar desert, Gujarat, Rajashtan
G.b. salinarum Salt range and E to Delhi
Naturally G&G elevate all subspecies to species status
Sample sizes
Sample sizes for skins are not given
Skulls/horns (Males/females)
bennettii 8-20/3-10
christyi 7-33/1-4
fuscifrons 15-22/6-11
salinarum 7-15/1-3
shikarii 3-4/1-2
Except for shikarii the other taxa have relatively ok sample sizes for males, but on the small side for multivariate analyses with 10 variables which was performed (exact results are not given). For females sample sizes range from extremely small to okish, but still too small for any multivariate analyses.
Skins
shikarii is reported as having a light skin colour, whereas fuscifrons is said to be dark, bennettii is said to be dull reddish brown, christyi very pale drab brown and salinarum a rich tobacco brown. salinarum also has no contrasting flank band, which is said to be prominent in fuscifrons, christyi has slightly darker median dorsal and lower flank zones, which are clearly more tawny and abrubtly darker in bennettii. No dorsal band presence/absence is described for shikarii, but based on pictures there is one present very slightly. A nose spot is described only for fuscifrons, though christyi, salinarum and bennettii also clearly have one based on pictures in Castello's field guide. Only shikarii has a less pronounced spot.
Skulls & horns
Female horns are described to be very long in shikarii, but there are only 2 datapoints which fall completely in the variation of fuscifrons. Horn divergence (which I assume corresponds with their horn span measurement) is said to be largest in shikarii, but though true on average the variation falls completely within the other taxa and 3 of the 4 other taxa have higher maximum spans in males, for females there are hardly any data points available here. fuscifrons is described as having a small skull, especially for females, but this is not reflected in the data at all. The horns are also said to be much less divergent, but the variation falls completely within the other taxa. bennettii females are said to have larger nasals, but though true on average, the overlap in measurements is huge with the other taxa. The reported less divergent horns in bennettii males compared to the span is also not reflected in the data. The reported differences for christyi are also not represented in the data. The same goes for the reported unique characteristics of salinarum, though the horns are indeed slightly less turned inside than the other taxa, except bennettii, but see picture of fuscifrons below and in Castello's field guide, which suggest that this characteristic is also there in fuscifrons.
Additional data
I am not aware of any research looking at the genetical differences between the reported subspecies.
Summarizing
Overall I see no reason to recognize any of the proposed splits. There are absolutely no clear differences in the skull and horn measurements and though there are some slight differences between the skin, especially in overall colour and how prominent the flank band is, that is far too little to warrant species status.
Gazella bennettii (likely bennettii)
@Giant Eland , New Delhi Zoo, India
Gazella bennettii (likely fuscifrons)
@J I N X , Peshawar Zoo, Pakistan
Gazella bennettii shikarii
@fofo , Iran
Gazella bennetti christyi
@toto98 , Thar Desert, India
No pictures of salinarum have been uploaded yet to the gallery.
next: the remaining Antilopini.
Blackbuck
The Blackbuck (Antilope cervicapra) is an antelope from the Indian peninsula. Currently two subspecies are recognized:
A.c. cervicapra E and S of the Delhi Region, India
A.c. rajputanae W of the Delhi Region into Pakistan
Antilope cervicapra rajputanae
@Chlidonias , Kanjari Blackbuck Reserve, India
Blackbuck gazelle female:
@Azubaa , ZSL Whipsnade Zoo, UK
No pictures of the nominate cervicapra have been uploaded to the gallery.
Tibetan gazelle
The Tibetan gazelle (Procabra picticaudata) is an antelope from the Tibetan plateau. It is currently considered monotypic.
females:
@baboon , Silling Lake, China
Przewalski's gazelle
The Przewalski's gazelle (Procapra przewalskii) is closely related to the slightly smaller Tibetan gazelle and has a limited distribution around Qinghai Lake in China. G&G recognize two subspecies diversicornis and przewalskii, this is not followed by HMW, as their relationship is not clear.
@Deer Forest , Qinghai-Tibet Plateau Wildlife Zoo, China
Mongolian gazelle
The Mongolian gazelle (Procapra gutturosa) is the larger cousin of the Tibetan and Przewalski's gazelle. This species is particularly known for the huge migrations it makes and is still one of the most numerous ungulates worldwide, despite heavy hunting pressure.
@Sun Wukong , Beijing Zoo, China
Sand gazelle
The Sand gazelle (Gazella marica) was formerly considered a subspecies of the Goitered gazelle, but genetic research has shown it to be more closely related to the Slender-horned and Cuvier's gazelle.
@Oryx , Sjarjah Desert Park, UAE
Mountain gazelle
The Mountain gazelle (Gazella gazella) was formerly considered conspecific with the Arabian gazelle, but has been shown to be clearly genetically and morphologically distinct and is now treated as a monotypic species.
@alexkant , Hai Park, Israel
Saudi gazelle
The Saudi gazelle (Gazella saudiya) is an extinct species of gazelle from the Arabian peninsula. It has been treated as a subspecies of the Dorcas gazelle, but genetic research has indicated it could represent a distinct species. Though this species is reportedly present in captivity, these populations are hybrids.
No pictures of the Saudi gazelle have been uploaded.
next: Takin.
The Blackbuck (Antilope cervicapra) is an antelope from the Indian peninsula. Currently two subspecies are recognized:
A.c. cervicapra E and S of the Delhi Region, India
A.c. rajputanae W of the Delhi Region into Pakistan
Antilope cervicapra rajputanae
@Chlidonias , Kanjari Blackbuck Reserve, India
Blackbuck gazelle female:
@Azubaa , ZSL Whipsnade Zoo, UK
No pictures of the nominate cervicapra have been uploaded to the gallery.
Tibetan gazelle
The Tibetan gazelle (Procabra picticaudata) is an antelope from the Tibetan plateau. It is currently considered monotypic.
females:
@baboon , Silling Lake, China
Przewalski's gazelle
The Przewalski's gazelle (Procapra przewalskii) is closely related to the slightly smaller Tibetan gazelle and has a limited distribution around Qinghai Lake in China. G&G recognize two subspecies diversicornis and przewalskii, this is not followed by HMW, as their relationship is not clear.
@Deer Forest , Qinghai-Tibet Plateau Wildlife Zoo, China
Mongolian gazelle
The Mongolian gazelle (Procapra gutturosa) is the larger cousin of the Tibetan and Przewalski's gazelle. This species is particularly known for the huge migrations it makes and is still one of the most numerous ungulates worldwide, despite heavy hunting pressure.
@Sun Wukong , Beijing Zoo, China
Sand gazelle
The Sand gazelle (Gazella marica) was formerly considered a subspecies of the Goitered gazelle, but genetic research has shown it to be more closely related to the Slender-horned and Cuvier's gazelle.
@Oryx , Sjarjah Desert Park, UAE
Mountain gazelle
The Mountain gazelle (Gazella gazella) was formerly considered conspecific with the Arabian gazelle, but has been shown to be clearly genetically and morphologically distinct and is now treated as a monotypic species.
@alexkant , Hai Park, Israel
Saudi gazelle
The Saudi gazelle (Gazella saudiya) is an extinct species of gazelle from the Arabian peninsula. It has been treated as a subspecies of the Dorcas gazelle, but genetic research has indicated it could represent a distinct species. Though this species is reportedly present in captivity, these populations are hybrids.
No pictures of the Saudi gazelle have been uploaded.
next: Takin.
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Any idea on subspecies of Blackbuck in captivity? I've heard rajputanae for Bronx's but I don't know how accurate that is.
~Thylo
~Thylo
Morphologically most Blackbuck pictures here resemble rajputanae (amount of white on legs and large amount of white on the sides). Castello also has rajputanae pictures included which were taken in European and American zoos. So it could well be that the imports in both Europe and US were pure rajputanae, but it would be somewhat surprising if the nominate never made it to Europe...
Takin
The Takin (Budorcas taxicolor) is a large caprine that has long been thought to be most closely related to the Musk ox. The Takin is however most closely related to the Rocky Mountain goat and Ibexes, whereas the Musk ox is most closely related to the Gorals. Traditionally four Takin subspecies have been recognized:
B.t. taxicolor, Mishmi takin, N Myanmar and Yunnan, China,
B.t. whitei, Bhutan takin, Bhutan and SE Tibet
B.t. tibetanus, Sichuan takin, Sichuan, China
B.t. bedfordi, Golden takin, Qinling Range, China
G&G elevate all subspecies to species status.
Sample sizes
Sample sizes for skins are not reported, though it is noted that pelage characteristics are also based on living Takin in Tierpark Berlin and the Shanghai Zoo breeding farm.
Horns/skulls (males/females)
bedfordi 1-3/2-5
taxicolor 3-12/1-5
tibetanus 3-16/0-5
whitei 2-8/0-2
Sample sizes are extremely limited for all taxa in the vast majority of measurements taken.
Skins
whitei and taxicolor are extremely similar in pelage color, though whitei is described as somewhat darker according to Castello. G&G describe whitei has having the same pelage in both juveniles and adults as juvenile taxicolor. Where in taxicolor the blackish colour extends to the whole face, under the neck and to the lower flanks. taxicolor develops a pale yellow tone on the upper parts. bedfordi is described as young animals being light golden-brown, with dark only on the haunches and a wholly light-toned face. With age the colour in bedfordi becomes more red-gold tone and develops a pale golden saddle. But the face nevers becomes dark. tibetanus young animals are golden-brown with much less extensive dark then taxicolor, but animals become darker with age, with a conspicious pale saddle and with age clearly black on the nose.
Skulls/horns
whitei is distinguished by having relatively large teeth, but there is almost complete overlap with the other taxa. whitei is described as having small horns with a narrow span, but there is complete overlap with tibetanus in horn span and horn length is not given in the table. whitei males are described as being small, but overlap widely in measurements with the other taxa. whitei females have somewhat smaller horns (but n=2) than males, but are similar in size. Based on the extremely limited dataset taxicolor and tibetanus are clearly sexually dimorphic with larger males, bedfordi seems less sexually dimorphic. In all three taxa male horns have larger spans and tip-to-tip distances. On average the nasal region is less inflated in bedfordi and more in tibetanus, though true on average there is some overlap and sample sizes are extremely small. In overall size bedfordi and tibetanus are somewhat larger than taxicolor and whitei, but sample sizes are very small and there is overlap in the measurements.
Additional data
G&G cite Li et al. (2003) to show that there are clear genetic differences between bedfordi, tibetanus and taxicolor, no samples of whitei were taken. Li et al. (2003) do however have only 1 taxicolor sample. Additionally the genetic differences are rather small, Genetic divergence is maximum 2.5% between the subspecies. And the difference between bedfordi and tibetanus is much smaller and more around 1%.
Summarizing
Though there appear to be some differences in pelage and sexual dimorphism, which is something that visitors to Tierpark Berlin can attest to, the differences are overall rather minor. This is especially true when considering the extremely small sample size and thus the reduced possibilities to draw any hard conclusions. The genetics also contradict a split as though there are genetic differences between the taxa, these differences are very small, especially compared to other ungulates. I therefore see no reason to elevate any of the subspecies to species status, it even seems questionable to me whether whitei should be treated as a valid subspecies or whether it should be resumed into taxicolor, but more morphological and genetic data is needed for that.
Budorcas taxicolor taxicolor
Male:
@gentle lemur , Paignton Zoo, UK
female:
@bongowwf , Zoo Antwerpen, Belgium
Budorcas taxicolor whitei
@Shirokuma , Motithan Takin Preserve, Bhutan
Budorcas taxicolor bedfordi
@Michal Sloviak , Bojnice Zoo, Slovakia
Budorcas taxicolor tibetanus
@Deer Forest , Tangjiahe National Nature Reserve, China
Reference
Li et al. (2003): An Error Occurred Setting Your User Cookie
Next: Pyrenean chamois
The Takin (Budorcas taxicolor) is a large caprine that has long been thought to be most closely related to the Musk ox. The Takin is however most closely related to the Rocky Mountain goat and Ibexes, whereas the Musk ox is most closely related to the Gorals. Traditionally four Takin subspecies have been recognized:
B.t. taxicolor, Mishmi takin, N Myanmar and Yunnan, China,
B.t. whitei, Bhutan takin, Bhutan and SE Tibet
B.t. tibetanus, Sichuan takin, Sichuan, China
B.t. bedfordi, Golden takin, Qinling Range, China
G&G elevate all subspecies to species status.
Sample sizes
Sample sizes for skins are not reported, though it is noted that pelage characteristics are also based on living Takin in Tierpark Berlin and the Shanghai Zoo breeding farm.
Horns/skulls (males/females)
bedfordi 1-3/2-5
taxicolor 3-12/1-5
tibetanus 3-16/0-5
whitei 2-8/0-2
Sample sizes are extremely limited for all taxa in the vast majority of measurements taken.
Skins
whitei and taxicolor are extremely similar in pelage color, though whitei is described as somewhat darker according to Castello. G&G describe whitei has having the same pelage in both juveniles and adults as juvenile taxicolor. Where in taxicolor the blackish colour extends to the whole face, under the neck and to the lower flanks. taxicolor develops a pale yellow tone on the upper parts. bedfordi is described as young animals being light golden-brown, with dark only on the haunches and a wholly light-toned face. With age the colour in bedfordi becomes more red-gold tone and develops a pale golden saddle. But the face nevers becomes dark. tibetanus young animals are golden-brown with much less extensive dark then taxicolor, but animals become darker with age, with a conspicious pale saddle and with age clearly black on the nose.
Skulls/horns
whitei is distinguished by having relatively large teeth, but there is almost complete overlap with the other taxa. whitei is described as having small horns with a narrow span, but there is complete overlap with tibetanus in horn span and horn length is not given in the table. whitei males are described as being small, but overlap widely in measurements with the other taxa. whitei females have somewhat smaller horns (but n=2) than males, but are similar in size. Based on the extremely limited dataset taxicolor and tibetanus are clearly sexually dimorphic with larger males, bedfordi seems less sexually dimorphic. In all three taxa male horns have larger spans and tip-to-tip distances. On average the nasal region is less inflated in bedfordi and more in tibetanus, though true on average there is some overlap and sample sizes are extremely small. In overall size bedfordi and tibetanus are somewhat larger than taxicolor and whitei, but sample sizes are very small and there is overlap in the measurements.
Additional data
G&G cite Li et al. (2003) to show that there are clear genetic differences between bedfordi, tibetanus and taxicolor, no samples of whitei were taken. Li et al. (2003) do however have only 1 taxicolor sample. Additionally the genetic differences are rather small, Genetic divergence is maximum 2.5% between the subspecies. And the difference between bedfordi and tibetanus is much smaller and more around 1%.
Summarizing
Though there appear to be some differences in pelage and sexual dimorphism, which is something that visitors to Tierpark Berlin can attest to, the differences are overall rather minor. This is especially true when considering the extremely small sample size and thus the reduced possibilities to draw any hard conclusions. The genetics also contradict a split as though there are genetic differences between the taxa, these differences are very small, especially compared to other ungulates. I therefore see no reason to elevate any of the subspecies to species status, it even seems questionable to me whether whitei should be treated as a valid subspecies or whether it should be resumed into taxicolor, but more morphological and genetic data is needed for that.
Budorcas taxicolor taxicolor
Male:
@gentle lemur , Paignton Zoo, UK
female:
@bongowwf , Zoo Antwerpen, Belgium
Budorcas taxicolor whitei
@Shirokuma , Motithan Takin Preserve, Bhutan
Budorcas taxicolor bedfordi
@Michal Sloviak , Bojnice Zoo, Slovakia
Budorcas taxicolor tibetanus
@Deer Forest , Tangjiahe National Nature Reserve, China
Reference
Li et al. (2003): An Error Occurred Setting Your User Cookie
Next: Pyrenean chamois
Chamois
Most authors have recognized two species of Chamois: the Pyrenean chamois (Rupicapra pyrenaica) of Spain and central Italy and the Alpine chamois (Rupicapra rupicapra) of Central and Eastern Europe and Asia Minor and the Caucasus. The Chamois case is a very interesting one and even though the Pyrenean and the Alpine chamois are clearly seperate species based on behaviour, genetics and morphology the relationships among multiple Chamois populations are often rather unclear and there are natural hybrids. Therefore I will treat both species in the same account.
Traditionally the Pyrenaen chamois is subdivided into three subspecies
R.p. pyrenaica Pyrenees ,on the border of France and Spain, including Andorra
R.p. parva Cantabrian Mountains in NW Spain
R.p. ornata Abruzzi Mountains in Central Italy
The Alpine chamois has traditionally been divided in 7 subspecies
R.r. rupicapra Alps, Jura Mountains, Vogues, Black Forest
R.r. tatrica Tatra mountains on the border of Slovakia and Poland
R.r. balcanica Balkans from Croatia to Greece and Bulgaria
R.r. carpatica Carpathian Mountains in Romania
R.r. cartusiana Chartreuse Mountains in SE France
R.r. asiatica Turkey and SW Georgia
R.r. caucasica Caucasus Mountains in Azerbaijan, Georgia & Russia
For a map see Figure 1 in Rodriquez et al. (2010) in the reference list.
G&G elevate most subspecies to species status, except that they lump caucasica and asiatica and they treat cartusiana as subspecies of rupicapra. Additionally they are unsure of the position of R.r. balcanica and in HMW this taxon is treated as a subspecies of rupicapra.
Sample size
No sample sizes for skins are given, though it appears they have seen 0 skins of caucasica or asiatica, nor are skin characteristics given for ornata, parva, carpatica or balcanica
Skulls (males/females)
cartusiana 8-9/5
rupicapra 6-7/10-11
tatrica 0-1/0-1
balcanica 2-6/1-5
asiatica 0-3/0-1
caucasica 0-4/0-2
carpatica 2-3/1
parva 0-1/0
pyrenaica 4/10-11
ornata 0-2/0-3
Horns (males/females)
cartusiana 8/5
rupicapra 14/18
tatrica 0/0
balcanica 2-4/1
asiatica 0-3/0-1
caucasica 4/2
carpatica 2/1
parva 0-1/0
pyrenaica 3/8-9
ornata 0-1/0-2
Sample sizes are thus extremely small for most taxa and even for the most sampled taxa rupicapra, cartusiana and pyrenaica sample sizes are rather low.
Additionally G&G cite a paper by Scala & Lovari (1984) that compares horns and skulls of ornata with pyrenaica, who sampled 9 males of each taxon for skulls and 13 pyrenaica and 18 ornata for skull measurements.
Finally G&G also cite an earlier paper by Scala & Lovari (1980) on skull and horn characteristics of carpatica (sample size = 5), balcanica (7), caucasica (6) and cartusiana (15). G&G partly base their assessment on them
Skins
As G&G do not give any details on the skins of most taxa, the following descriptions are taken from Castello's Bovid field guide. pyrenaica is described as darker than parva. ornata has a distinct white throat patch and large white areas on the side and back than the other taxa. tatrica is described as similar to rupicapra, whereas balcanica is said to be somewhat lighter than rupicapra. carpathica is described as darker than the other taxa. asiatica and caucasica are described as darker than rupicapra. caucasica is said to differ from carpathica by having a lighter coloration around the base of the horns.
Skulls and horns
As G&G had an extremely low sample size for ornata and also a low one for pyrenaica for that comparison I will use the data by Scala & Lovari (1984). They found no clear difference between the horns of ornata and pyrenaica. Both taxa separated well however based on skull measurements, ornata being characterized by a its great fontanelle length. parva is said to be smaller than pyrenaica and based on 2 skull measurements of one parva skull that seems to be true, though the third measurement (teeth length) overlaps. balcanica and cartusiana samples fall well within the variation of rupicapra. G&G use the data by Scala & Lovari (1980) to argue for the differences between rupicapra (cartusiana), carpatica and caucasica. They state that carpatica is larger than the other taxa and though true on average there is overlap in the measurements with both balcanica and cartusiana. Though caucasica is said to differ based on multivariate analysis, sample sizes are inadequate for that method and there is wide overlap with the other taxa when looking at single measurements. G&G cite Hrabe & Koubek (1984) to state that tatrica has narrower nasals in both sexes compared with other Chamois and the males also have shorter nasals. As this publication seems unavailable online I can't say anything on its accuracy.
Additional data
In addition to the morphological research cited above, there has been considerable genetic work on Chamois, most notably by Rodriquez et al. (2010). They used a combination of methods, analysing both mitochondrial DNA as well as multiple microsatellites (nuclear DNA). To make a long and complex story short, they find support for the genetic separation of all traditionally recognized taxa. But that is where it starts to get complext. both the Chamois from the SW Alps and the cartusiana Chamois are actually ancient hybrids between Pyrenean and Alpine chamois. Because of multiple Ice Ages, ranges of the Chamois, a mountain species have expanded and contracted considerably in the past, resulting in some ancient hybrid populations and isolated subspecies throughout Europe and W Asia. Apart from these two hybrid populations, only ornata from the Abruzzi Mountains in C Italy stands out as really distinct, forming one of the 3 mitochondrial clades (together with the hybrid cartusiana chamois). The other mitochondrial clades are pyrenaica + parva and rupicapra + balcanica + tatrica + carpatica + caucasica + asiatica. The animals of the SW Alps where of mixed mitochondrial origin. Using nuclear dna microsatellites cartusiana grouped strongly with rupicapra indicating its hybrid origin.
Summarizing
There does not seem to be much evidence for recognizing 6+ Chamois species, though through a very strict PSC approach one would be able to do so.... Combining morphological with genetical analyses there does however seem to be a case to elevate ornata to species status. This would mean that we would recognize three Chamois species: pyrenaica (pyrenaica + parva), ornata and rupicapra (rupicapra + balcanica + tatrica + carpatica + caucasica + asiatica). cartusiana could be treated as a unique hybrid (species?) between rupicapra and ornata. Such a situation is similar to the Red wolf in the US. For sure the last word has not been said yet about this complex species complex. What is clear however is that G&G's treatment is an oversimplified and in the case of cartusiana wrong assessment of the diversity in the genus Rupicapra.
References
Lovari & Scala (1980): https://www.tandfonline.com/doi/pdf/10.1080/11250008009440328
Rodriquez et al. (2010): Integrating phylogeographic patterns of microsatellite and mtDNA divergence to infer the evolutionary history of chamois (genus Rupicapra) | BMC Evolutionary Biology | Full Text
Scala & Lovari (1984): https://www.tandfonline.com/doi/pdf/10.1080/11250008409439467
Pyrenean chamois
Rupicapra pyrenaica pyrenaica
@Maguari , Domaine de Pescheray, France
Rupicapra pyrenaica parva
@Maguari , Picos de Europa NP, Spain
Abruzzi chamois
Rupicapra (pyrenaica) ornata
@Eagle, Tierpark Hellabrunn, Muenchen, Germany
Alpine chamois
Rupicapra rupicapra rupicapra
Winter pelage
@Tarsius, Tiergarten Hellbrunn, Salzburg, Austria
Summer pelage
@gentle lemur , Blackpool Zoo, UK
No pictures of balcanica, tatrica, carpatica, caucasica or asiatica have been uploaded to the gallery
next: Himalayan goral
Most authors have recognized two species of Chamois: the Pyrenean chamois (Rupicapra pyrenaica) of Spain and central Italy and the Alpine chamois (Rupicapra rupicapra) of Central and Eastern Europe and Asia Minor and the Caucasus. The Chamois case is a very interesting one and even though the Pyrenean and the Alpine chamois are clearly seperate species based on behaviour, genetics and morphology the relationships among multiple Chamois populations are often rather unclear and there are natural hybrids. Therefore I will treat both species in the same account.
Traditionally the Pyrenaen chamois is subdivided into three subspecies
R.p. pyrenaica Pyrenees ,on the border of France and Spain, including Andorra
R.p. parva Cantabrian Mountains in NW Spain
R.p. ornata Abruzzi Mountains in Central Italy
The Alpine chamois has traditionally been divided in 7 subspecies
R.r. rupicapra Alps, Jura Mountains, Vogues, Black Forest
R.r. tatrica Tatra mountains on the border of Slovakia and Poland
R.r. balcanica Balkans from Croatia to Greece and Bulgaria
R.r. carpatica Carpathian Mountains in Romania
R.r. cartusiana Chartreuse Mountains in SE France
R.r. asiatica Turkey and SW Georgia
R.r. caucasica Caucasus Mountains in Azerbaijan, Georgia & Russia
For a map see Figure 1 in Rodriquez et al. (2010) in the reference list.
G&G elevate most subspecies to species status, except that they lump caucasica and asiatica and they treat cartusiana as subspecies of rupicapra. Additionally they are unsure of the position of R.r. balcanica and in HMW this taxon is treated as a subspecies of rupicapra.
Sample size
No sample sizes for skins are given, though it appears they have seen 0 skins of caucasica or asiatica, nor are skin characteristics given for ornata, parva, carpatica or balcanica
Skulls (males/females)
cartusiana 8-9/5
rupicapra 6-7/10-11
tatrica 0-1/0-1
balcanica 2-6/1-5
asiatica 0-3/0-1
caucasica 0-4/0-2
carpatica 2-3/1
parva 0-1/0
pyrenaica 4/10-11
ornata 0-2/0-3
Horns (males/females)
cartusiana 8/5
rupicapra 14/18
tatrica 0/0
balcanica 2-4/1
asiatica 0-3/0-1
caucasica 4/2
carpatica 2/1
parva 0-1/0
pyrenaica 3/8-9
ornata 0-1/0-2
Sample sizes are thus extremely small for most taxa and even for the most sampled taxa rupicapra, cartusiana and pyrenaica sample sizes are rather low.
Additionally G&G cite a paper by Scala & Lovari (1984) that compares horns and skulls of ornata with pyrenaica, who sampled 9 males of each taxon for skulls and 13 pyrenaica and 18 ornata for skull measurements.
Finally G&G also cite an earlier paper by Scala & Lovari (1980) on skull and horn characteristics of carpatica (sample size = 5), balcanica (7), caucasica (6) and cartusiana (15). G&G partly base their assessment on them
Skins
As G&G do not give any details on the skins of most taxa, the following descriptions are taken from Castello's Bovid field guide. pyrenaica is described as darker than parva. ornata has a distinct white throat patch and large white areas on the side and back than the other taxa. tatrica is described as similar to rupicapra, whereas balcanica is said to be somewhat lighter than rupicapra. carpathica is described as darker than the other taxa. asiatica and caucasica are described as darker than rupicapra. caucasica is said to differ from carpathica by having a lighter coloration around the base of the horns.
Skulls and horns
As G&G had an extremely low sample size for ornata and also a low one for pyrenaica for that comparison I will use the data by Scala & Lovari (1984). They found no clear difference between the horns of ornata and pyrenaica. Both taxa separated well however based on skull measurements, ornata being characterized by a its great fontanelle length. parva is said to be smaller than pyrenaica and based on 2 skull measurements of one parva skull that seems to be true, though the third measurement (teeth length) overlaps. balcanica and cartusiana samples fall well within the variation of rupicapra. G&G use the data by Scala & Lovari (1980) to argue for the differences between rupicapra (cartusiana), carpatica and caucasica. They state that carpatica is larger than the other taxa and though true on average there is overlap in the measurements with both balcanica and cartusiana. Though caucasica is said to differ based on multivariate analysis, sample sizes are inadequate for that method and there is wide overlap with the other taxa when looking at single measurements. G&G cite Hrabe & Koubek (1984) to state that tatrica has narrower nasals in both sexes compared with other Chamois and the males also have shorter nasals. As this publication seems unavailable online I can't say anything on its accuracy.
Additional data
In addition to the morphological research cited above, there has been considerable genetic work on Chamois, most notably by Rodriquez et al. (2010). They used a combination of methods, analysing both mitochondrial DNA as well as multiple microsatellites (nuclear DNA). To make a long and complex story short, they find support for the genetic separation of all traditionally recognized taxa. But that is where it starts to get complext. both the Chamois from the SW Alps and the cartusiana Chamois are actually ancient hybrids between Pyrenean and Alpine chamois. Because of multiple Ice Ages, ranges of the Chamois, a mountain species have expanded and contracted considerably in the past, resulting in some ancient hybrid populations and isolated subspecies throughout Europe and W Asia. Apart from these two hybrid populations, only ornata from the Abruzzi Mountains in C Italy stands out as really distinct, forming one of the 3 mitochondrial clades (together with the hybrid cartusiana chamois). The other mitochondrial clades are pyrenaica + parva and rupicapra + balcanica + tatrica + carpatica + caucasica + asiatica. The animals of the SW Alps where of mixed mitochondrial origin. Using nuclear dna microsatellites cartusiana grouped strongly with rupicapra indicating its hybrid origin.
Summarizing
There does not seem to be much evidence for recognizing 6+ Chamois species, though through a very strict PSC approach one would be able to do so.... Combining morphological with genetical analyses there does however seem to be a case to elevate ornata to species status. This would mean that we would recognize three Chamois species: pyrenaica (pyrenaica + parva), ornata and rupicapra (rupicapra + balcanica + tatrica + carpatica + caucasica + asiatica). cartusiana could be treated as a unique hybrid (species?) between rupicapra and ornata. Such a situation is similar to the Red wolf in the US. For sure the last word has not been said yet about this complex species complex. What is clear however is that G&G's treatment is an oversimplified and in the case of cartusiana wrong assessment of the diversity in the genus Rupicapra.
References
Lovari & Scala (1980): https://www.tandfonline.com/doi/pdf/10.1080/11250008009440328
Rodriquez et al. (2010): Integrating phylogeographic patterns of microsatellite and mtDNA divergence to infer the evolutionary history of chamois (genus Rupicapra) | BMC Evolutionary Biology | Full Text
Scala & Lovari (1984): https://www.tandfonline.com/doi/pdf/10.1080/11250008409439467
Pyrenean chamois
Rupicapra pyrenaica pyrenaica
@Maguari , Domaine de Pescheray, France
Rupicapra pyrenaica parva
@Maguari , Picos de Europa NP, Spain
Abruzzi chamois
Rupicapra (pyrenaica) ornata
@Eagle, Tierpark Hellabrunn, Muenchen, Germany
Alpine chamois
Rupicapra rupicapra rupicapra
Winter pelage
@Tarsius, Tiergarten Hellbrunn, Salzburg, Austria
Summer pelage
@gentle lemur , Blackpool Zoo, UK
No pictures of balcanica, tatrica, carpatica, caucasica or asiatica have been uploaded to the gallery
next: Himalayan goral
Himalayan goral
The Himalayan goral (Naemorhedus goral*) is a small caprine that occurs on the lower slopes of the Himalayas between 900 and 4000 meters altitude. It occurs from Pakistan to Eastern India, including Nepal and Bhutan and extreme S China. The Himalayan goral is sometimes considered monotypic or alternatively two subspecies are recognized:
N.g. goral (Himalayan brown goral) E of the Sutlej river, from Himachal Pradesh, India, east through Nepal, China and Bhutan
N.g. bedfordi (Himalayan gray goral) W of the Sutlej river, from Himachal Pradesh, India, west to Kashmir and Pakistan
G&G elevate both subspecies to species status
Sample sizes
Skins
goral 21
bedfordi 13
Additionally about 10 live specimens of goral were seen at Delhi Zoo.
No sample sizes for skulls/horns are given
Skins
goral is described as being either medium brown, with black tips to the hairs or slightly grayer, or pale or dark fawn. The legs are browner to a very bright tan or white on the forelegs only. The underside is paler gray and throat and chin are described as variable white. Bedfordi is described as being gray-brown to yellow-gray with lighter and yellower legs, which have a dark brown line down the front. The underside is described as off-white and the chin is creamy with this tone reaching down the throat.
Skulls + horns
G&G indicate they did not find any differences in skulls or horns between both taxa, even though data for this are not presented. Castello states that bedfordi has more curved horns with heavier rings, but based on the bedfordi picture below there seems to be more variation than he describes.
Additional data
I am not aware of any genetical research on the differences between Himalayan goral populations
Summarizing
Giving the lack of skull and horn differences and the lack of genetic data, I don’t think that the difference in pelage colour warrant the recognition of two Himalayan goral species. The differences are however clear enough that the recognition of two subspecies is certainly warranted.
* Note that there is inconsistency about the exact spelling of Naemorhedus, with Nemorhaedus also being commonly used, but Naemorhedus is the original spelling, thus followed here
Naemorhedus goral bedfordi
@J I N X , Lalazar Wildlife Park, Pakistan
No pictures of N.g. goral have been uploaded to the gallery yet.
The Himalayan goral (Naemorhedus goral*) is a small caprine that occurs on the lower slopes of the Himalayas between 900 and 4000 meters altitude. It occurs from Pakistan to Eastern India, including Nepal and Bhutan and extreme S China. The Himalayan goral is sometimes considered monotypic or alternatively two subspecies are recognized:
N.g. goral (Himalayan brown goral) E of the Sutlej river, from Himachal Pradesh, India, east through Nepal, China and Bhutan
N.g. bedfordi (Himalayan gray goral) W of the Sutlej river, from Himachal Pradesh, India, west to Kashmir and Pakistan
G&G elevate both subspecies to species status
Sample sizes
Skins
goral 21
bedfordi 13
Additionally about 10 live specimens of goral were seen at Delhi Zoo.
No sample sizes for skulls/horns are given
Skins
goral is described as being either medium brown, with black tips to the hairs or slightly grayer, or pale or dark fawn. The legs are browner to a very bright tan or white on the forelegs only. The underside is paler gray and throat and chin are described as variable white. Bedfordi is described as being gray-brown to yellow-gray with lighter and yellower legs, which have a dark brown line down the front. The underside is described as off-white and the chin is creamy with this tone reaching down the throat.
Skulls + horns
G&G indicate they did not find any differences in skulls or horns between both taxa, even though data for this are not presented. Castello states that bedfordi has more curved horns with heavier rings, but based on the bedfordi picture below there seems to be more variation than he describes.
Additional data
I am not aware of any genetical research on the differences between Himalayan goral populations
Summarizing
Giving the lack of skull and horn differences and the lack of genetic data, I don’t think that the difference in pelage colour warrant the recognition of two Himalayan goral species. The differences are however clear enough that the recognition of two subspecies is certainly warranted.
* Note that there is inconsistency about the exact spelling of Naemorhedus, with Nemorhaedus also being commonly used, but Naemorhedus is the original spelling, thus followed here
Naemorhedus goral bedfordi
@J I N X , Lalazar Wildlife Park, Pakistan
No pictures of N.g. goral have been uploaded to the gallery yet.
Chinese goral
The Chinese goral (Naemorhedus griseus) is a small caprine that occurs in C and SE China as well as in Myanmar (Burma) and Thailand. Generally only two subspecies are recognized, though more have been described in the past, including arnouxianus which is how the Chinese goral in European zoos are treated.
N.g. griseus (Chinese goral) C and SE China, includes arnouxianus
N.g. evansi (Burmese goral) parts of Yunnan, S China, Myanmar (Burma) and NW Thailand
G&G elevate both subspecies to species status
Sample sizes
Skins
griseus 27
evansi 24
Additionally about 20 live griseus were observed in Chinese zoos.
No sample sizes for skulls/horns are given
Skins
griseus is described as being dark to light gray, with varying amounts of brown and a varying black overlay. Front and innerside of the shanks are very pale, whitish in juveniles, yellowish in adults. The thigh color is said to send a wedge down the outer side to halfway down the legs. legs often have a dark stripe down the front griseus hairs are fairly long and coars in all seasons. evansi is described as very light, fawn of somewhat brown (though pictures show very gray animals). The hair is said to be short in the available specimens, but seasonal variation can’t be excluded. The legs are mostly golden to creamy gold, with a darker wedge or browner, with a thick black line down the front. Though the throat for both species is described as white with a golden tone, bedfordi throats seem much whiter on the available pictures and the more golden throat of many griseus specimens
Skulls + horns
Though none of the data are actually presented evansi is being described as much smaller than griseus and the horns are described as being much smaller as well, with evansi being similar in horn length as the Red goral (H. baileyi) and griseus being similar in horn length to the Himalayan goral (H. goral).
Additional data
A study using the whole mitochondrial genome by Hassanin et al. (2012) found that evansi is actually sister to the Red goral and should thus probably be treated as a separate species.
Summarizing
Though it is impossible to assess the data quality for the horn and skull measurements they seem to indicate a clear difference between evansi and griseus. The fact that in terms of size evansi is actually much closer to the Red goral is supported by the fact that the genetic data indicate that these are actually sister taxa. It thus seems valid to recognize evansi as a separate Goral species.
Naemorhedus (griseus) griseus
@Deer Forest , Beijing Zoo, China
Naemorhedus (griseus) evansi
@Chlidonias , Don Inthanon NP, Thailand
Reference
Hassanin et al. (2012): https://www.sciencedirect.com/science/article/pii/S1631069111002800
Next: Chinese serow
The Chinese goral (Naemorhedus griseus) is a small caprine that occurs in C and SE China as well as in Myanmar (Burma) and Thailand. Generally only two subspecies are recognized, though more have been described in the past, including arnouxianus which is how the Chinese goral in European zoos are treated.
N.g. griseus (Chinese goral) C and SE China, includes arnouxianus
N.g. evansi (Burmese goral) parts of Yunnan, S China, Myanmar (Burma) and NW Thailand
G&G elevate both subspecies to species status
Sample sizes
Skins
griseus 27
evansi 24
Additionally about 20 live griseus were observed in Chinese zoos.
No sample sizes for skulls/horns are given
Skins
griseus is described as being dark to light gray, with varying amounts of brown and a varying black overlay. Front and innerside of the shanks are very pale, whitish in juveniles, yellowish in adults. The thigh color is said to send a wedge down the outer side to halfway down the legs. legs often have a dark stripe down the front griseus hairs are fairly long and coars in all seasons. evansi is described as very light, fawn of somewhat brown (though pictures show very gray animals). The hair is said to be short in the available specimens, but seasonal variation can’t be excluded. The legs are mostly golden to creamy gold, with a darker wedge or browner, with a thick black line down the front. Though the throat for both species is described as white with a golden tone, bedfordi throats seem much whiter on the available pictures and the more golden throat of many griseus specimens
Skulls + horns
Though none of the data are actually presented evansi is being described as much smaller than griseus and the horns are described as being much smaller as well, with evansi being similar in horn length as the Red goral (H. baileyi) and griseus being similar in horn length to the Himalayan goral (H. goral).
Additional data
A study using the whole mitochondrial genome by Hassanin et al. (2012) found that evansi is actually sister to the Red goral and should thus probably be treated as a separate species.
Summarizing
Though it is impossible to assess the data quality for the horn and skull measurements they seem to indicate a clear difference between evansi and griseus. The fact that in terms of size evansi is actually much closer to the Red goral is supported by the fact that the genetic data indicate that these are actually sister taxa. It thus seems valid to recognize evansi as a separate Goral species.
Naemorhedus (griseus) griseus
@Deer Forest , Beijing Zoo, China
Naemorhedus (griseus) evansi
@Chlidonias , Don Inthanon NP, Thailand
Reference
Hassanin et al. (2012): https://www.sciencedirect.com/science/article/pii/S1631069111002800
Next: Chinese serow
So if one were to split N. griseus and evansi, would arnouxianus (also the taxa in the US) be a subspecies of the former or would the former remain a monotypic species?
~Thylo
~Thylo
That would be a junior synonym to griseus, which would remain a monotypic species. The review by G&G does not indicate any real difference within each taxon and my baseline is that when no difference is indicated in their review, there probably is no difference.
Chinese serow
The Chinese serow (Capricornis milneedwardsi) was previously considered a subspecies of the Sumatran serow (C. sumatraensis). This species was however split in 2005 in four species, presumably based mostly on morphology. Serow from Sumatra have a karyotype of 2n=46 and at least some of the mainland populations have a karyotype of 2n=48. It seems however unknown how karyotypes differ between all subspecies. Divergence between the different species seems relatively recent and has been estimated at 0.19-0.31 million years ago (Dou et al. 2016), which is similar to divergence between the different Hartebeest (sub)species. Traditionally Sumatran serow was only split in four, G&G do however split the Chinese serow in another two species:
C. milneedwardsi Central and Eastern China
C. maritimus Myanmar, Laos, Vietnam, Cambodia, Thailand
Sample sizes
Skins
milneedwardsi 50
maritimus 19
No sample sizes for skulls or horns are given.
Skins
milneedwardsi is described as having long lank hair, which is black and tending to reddish, especially on the flanks, rump and tail. maritimus is described as black or dark brown, with long white hair bases, thus looking gray or brindled and that mane is mixed with white or pale-yellow buff hairs. The mane in milneedwardsi is mainly silvery, though varying. milneedwardsi has a light underside (Though I can't clearly see this on pictures), no white nose, but a white upper-lip, with a broad white jaw-strak. The underside in maritimus is not white, often black, with white lips, often to the back of the eye and short white or golden-brown joaw-streaks. milneedwardsi has a smudgy lightthroat patch, which in maritimus is described as white or golden-brown hairs which sometimes form a big patch. maritimus legs are jet black on the upper half, reddish tan or creamy white on the lower half with a sharp division in between. milneedwardsi legs are described increasingly mingled with foxy red down the upper segments, shanks pure reddish with sometimes a sharp demarcation on the knee.
Skulls
milneedwardsi is described to sometimes have a sharp upper "shelf" to the pre-orbital fossa, lacking in maritimus. No skull measurements are given.
Additional data
G&G additionally present limited data on tail, hindfoot and ear length, but there is wide overlap in all measurements and very small sample sizes. Castello describes maritimus as having a distinctly lower shoulder height, but I have seen no data to back that up. I have not seen any genetic data comparing maritimus to milneedwards
Summarizing
Though there are some clear differences in pelage, the lack of evidence for any consistent differences in skulls or any genetic evidence, lead me to the conclusion that maritimus would be best categorized as a subspecies of milneedwardsi. From my point of view all Serow, excluding Taiwan and Japanese Serow, should be treated as a species group and research on genetics and skull characteristics in this species group would be highly valuable.
Capricornis milneedwardsi milneedwardsi
@YuanChang , Huangshan Wildlife Rescue Center, China
Capricornis milneedwardsi maritimus
@Giant Eland , Dusit Zoo, Thailand
Next Wild sheep
The Chinese serow (Capricornis milneedwardsi) was previously considered a subspecies of the Sumatran serow (C. sumatraensis). This species was however split in 2005 in four species, presumably based mostly on morphology. Serow from Sumatra have a karyotype of 2n=46 and at least some of the mainland populations have a karyotype of 2n=48. It seems however unknown how karyotypes differ between all subspecies. Divergence between the different species seems relatively recent and has been estimated at 0.19-0.31 million years ago (Dou et al. 2016), which is similar to divergence between the different Hartebeest (sub)species. Traditionally Sumatran serow was only split in four, G&G do however split the Chinese serow in another two species:
C. milneedwardsi Central and Eastern China
C. maritimus Myanmar, Laos, Vietnam, Cambodia, Thailand
Sample sizes
Skins
milneedwardsi 50
maritimus 19
No sample sizes for skulls or horns are given.
Skins
milneedwardsi is described as having long lank hair, which is black and tending to reddish, especially on the flanks, rump and tail. maritimus is described as black or dark brown, with long white hair bases, thus looking gray or brindled and that mane is mixed with white or pale-yellow buff hairs. The mane in milneedwardsi is mainly silvery, though varying. milneedwardsi has a light underside (Though I can't clearly see this on pictures), no white nose, but a white upper-lip, with a broad white jaw-strak. The underside in maritimus is not white, often black, with white lips, often to the back of the eye and short white or golden-brown joaw-streaks. milneedwardsi has a smudgy lightthroat patch, which in maritimus is described as white or golden-brown hairs which sometimes form a big patch. maritimus legs are jet black on the upper half, reddish tan or creamy white on the lower half with a sharp division in between. milneedwardsi legs are described increasingly mingled with foxy red down the upper segments, shanks pure reddish with sometimes a sharp demarcation on the knee.
Skulls
milneedwardsi is described to sometimes have a sharp upper "shelf" to the pre-orbital fossa, lacking in maritimus. No skull measurements are given.
Additional data
G&G additionally present limited data on tail, hindfoot and ear length, but there is wide overlap in all measurements and very small sample sizes. Castello describes maritimus as having a distinctly lower shoulder height, but I have seen no data to back that up. I have not seen any genetic data comparing maritimus to milneedwards
Summarizing
Though there are some clear differences in pelage, the lack of evidence for any consistent differences in skulls or any genetic evidence, lead me to the conclusion that maritimus would be best categorized as a subspecies of milneedwardsi. From my point of view all Serow, excluding Taiwan and Japanese Serow, should be treated as a species group and research on genetics and skull characteristics in this species group would be highly valuable.
Capricornis milneedwardsi milneedwardsi
@YuanChang , Huangshan Wildlife Rescue Center, China
Capricornis milneedwardsi maritimus
@Giant Eland , Dusit Zoo, Thailand
Next Wild sheep
Wild sheep
Taxonomy in the genus Ovis is among the most complicated of any bovid and between one and seven species have traditionally been recognized. In recent years Snow sheep (Ovis nivicola), Dall sheep (Ovis dalli) and Bighorn sheep (Ovis canadensis) have normally been treated as distinct species. The mess starts with when it comes to the sheep ranging from Europe to Central Asia. There has been a lot of confusion on the number of species recognized here, which has been made extra confusing by a large number of seemingly hybrid populations. Traditionally are the Argali (Ovis ammon), Urial (Ovis vignei) and the Asiatic mouflon (Ovis gmelinii) and the European mouflon (Ovis orientalis), where the last three have often been considered conspecific. Another problem is that orientalis has probably been described based on a hybrid. The Mouflon from Europe (musimon) are however descended of early-domesticated sheep returned to a wild life (e.g. Hiendleder et al. 2002), as are likely Cypriotic mouflon (ophion) (Sanna et al. 2016). These two taxa are not considered any further.
G&G recognize the following species within these species groups:
Ovis gmelini group (Ovis orientalis) = Mouflon
O. gmelini Turkey, Armenia, SW Azerbaijan, NE Iraq, NW Iran
O. isphaganica WC Iran, mountains around Esfahan city
O. laristanica Laristan and Fars province, Iran
Ovis vignei group = Urial
O. vignei, Ladakh, Kashmir in Pakistan and India and Wakhan and Ishkashim in Tajikistan
O. punjabiensis, Punjab province, Pakistan
O. bochariensis, NE Turkmenistan, S Uzbekistan, N Afghanistan, S Tajikistan
O. arabica, Oman and SE Saudi Arabia
O. cycloceros (including arkal) S Turkmenistan, NE Iran, Afghanistan
O. severtzovi Uzbekistan and the Turkestan range in Kyrgyzstan and Tajikistan
Ovis ammon group = Argali
O. nigrimontana Darya Karatau mountains, Kazakhstan
O. karelini, Tian-Shan mountains in SE Kazakhstan and E Kyrgyzstan, E to Urumqi in W China
O. collium Kazakhstan and NW China
O. polli E part of the Wakhan corridor, Afghanistan, Pamir plateau, Tajikistan, Pamir region and SE Kyrgyzstan into W China
O. hodgsoni, N India, N Nepal, N Bhutan, Tibet, NW Sichuan and SW Gansu and S Qinghai, China
O. ammon Altai mountains, W Mongolia, N Xinjang China, NE Kazakstan and Gorno-Alta and Tuvai republics in Siberian Russia
O. darwini S Mongolia, N China
O. jubata NE China
Sample sizes
Sample sizes for skins are not given
Skulls/horns (males only)
gmelini 3-7
isphaganica 1-8
laristanica 1-2
cycloceros 18-36
bochariensis 6-10
vignei 4-10
punjabiensis 1-6
arabica 0-1
severtzovi 0-5
nigrimontana 0-6
karelini 9-19
polli 8-31
collium 0-7
ammon 0-19
darwini 0-3
hodgsoni 2-16
The many 0 are due to the fact that the frontal arc was not measured in many specimens.
Data for jubata are taken from (Geist 1991) and are thus not shown in the main table.
Additionally there are samples from several hybrid populations (gmelini X arkal and cycloceros X laristanica)
Skins
The main difference between the vignei and gmelini group is that the gmelini group has a white saddle patch, which is absent in vignei. There are several minor pelage differences withing the gmelini group. The main difference between the vignei and ammon group is that the vignei group has a clear mane, absent in the ammon group.
Horns & skulls
vignei group
Both isphaganica and laristanica are said to have shorter teeth and thinner horns than gmelini, but there is wide overlap in teeth measurements and there is only 1 horn measurement for isphaganica and 2 for laristanica, so no reliable conclusion can be drawn
gmelini group
punjabiensis and bochariensis are very similar in skull characteristics and have on average a larger frontal arc than cycloceros, but there is overlap between bochariensis an cycloceros. The single arabica skull seems to have shorter teeth than all other taxa, the skull in general fits this group, though in terms of distribution a placement in the vignei group is more likely. in Skull heigth and skull breadth severtzovi seems somewhat intermediate between the gmelini and ammon group.
ammon group
ammon and hodgsoni are on average the largest taxa and this difference is absolute compared to some other taxa, though there are several intermediate taxa as well nigrimontana, darwini and collium are the smaller taxa. darwini is described as having a shorter, narrower but somewhat higher skull compared to hodgsoni and ammon, there is however large overlap in these measurements and sample sizes to small for reliable multivariate analyses. Overall a larger number of more subjective skull and horn measures are described, but given the data provided in the table combined with small sample sizes it is hard to draw generalizations from them.
Additional data
There has been quite some research on the genetics of wild sheep, especially because they are the forefathers of domestic sheep. There are clear differences genetically between the three groups. gmelini has a karyotype of 2n=54, vignei one of 2n=58 and ammon one of 2n=56. Research by Rezaei et al. (2010) indicate that ammon likely diverged from gmelini + vignei +- 1.72 million years ago (+-0.36 mya) and gmelini and vignei likely split 1.26 million years ago (+-0.36 mya). The relationships are however made more complex by extensive hybridization at the borders of these groups. severtzovi clearly groups in the ammon group. Several taxa: isphanicia, arkal, punjabiensis and gmelini are found not to be monophyletic based on genetic data.
Summarizing
Using a combination of morphological and genetic research it seems best to recognize three species of wild sheep in Eurasia next to the Snow sheep: Ovis gmelini (=Ovis orientalis) , Ovis vignei and Ovis ammon. Within each species there is quite some variation and all taxa elevated to species status by G&G can at least for now be recognized as subspecies, genetic research is however necessary to understand further the exact phylogenetic relationships within each group. Additionally the status of wild sheep on the Arabian peninsula is of conservation concern, as it is unclear whether they belong to vignei or gmelini.
Mouflon
Ovis gmelini gmelini
@Tomek , Tallinn Zoo, Estonia
Urial
Ovis vignei vignei
@Chlidonias , Hemis NP, India
Ovis vignei cycloceros (arkal)
@Nick@Amsterdam, Tierpark Berlin, Germany
Ovis vignei punjabiensis
@J I N X , Peshawar Zoo, Pakistan
Ovis vignei bochariensis
@Tomek , Zoo Liberec, Czechia
Argali
Ovis ammon ammon
@Giant Eland , Novosibirsk Zoo, Russia
Ovis ammon collium
@alexkant , Karaganda Zoo, Kazakhstan
Ovis ammon hodgsoni (including dalailamae)
@Deer Forest , Beijing Zoo
Ovis ammon jubata
@bongowwf , Shanghai Zoo, China
Ovis ammon polli
@Dormitator , Moscow Zoo breeding center, Sychovo, Russia
Ovis ammon darwini
@Chlidonias , Ikh Nart Nature Reserve, Mongolia
Hybrid sheep
Hybrid vignei X gmelini "Alborz sheep"
@fofo , Tehran Zoo, Iran
European mouflon
Ovis varies var. ophion Cyprus mouflon
@NigeW , Pafos Zoo, Cyprus
Ovis varies var. musimon European mouflon
@Tomek , New Forest Wildlife Park, UK
No pictures of nigrimontana, severtzovi, karelini, arabica, isphaganica or laristanica have been uploaded to the gallery yet
References
Hiendleder et al. (2002): https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1690972/pdf/12028771.pdf
Rezaei et al. (2010): https://www.sciencedirect.com/science/article/pii/S1055790309004461?via=ihubhttps://www.sciencedirect.com/science/article/pii/S1055790309004461?via=ihub
Sanna et al. (2016): The First Mitogenome of the Cyprus Mouflon (Ovis gmelini ophion): New Insights into the Phylogeny of the Genus Ovis
Next: the remaining Caprini
Taxonomy in the genus Ovis is among the most complicated of any bovid and between one and seven species have traditionally been recognized. In recent years Snow sheep (Ovis nivicola), Dall sheep (Ovis dalli) and Bighorn sheep (Ovis canadensis) have normally been treated as distinct species. The mess starts with when it comes to the sheep ranging from Europe to Central Asia. There has been a lot of confusion on the number of species recognized here, which has been made extra confusing by a large number of seemingly hybrid populations. Traditionally are the Argali (Ovis ammon), Urial (Ovis vignei) and the Asiatic mouflon (Ovis gmelinii) and the European mouflon (Ovis orientalis), where the last three have often been considered conspecific. Another problem is that orientalis has probably been described based on a hybrid. The Mouflon from Europe (musimon) are however descended of early-domesticated sheep returned to a wild life (e.g. Hiendleder et al. 2002), as are likely Cypriotic mouflon (ophion) (Sanna et al. 2016). These two taxa are not considered any further.
G&G recognize the following species within these species groups:
Ovis gmelini group (Ovis orientalis) = Mouflon
O. gmelini Turkey, Armenia, SW Azerbaijan, NE Iraq, NW Iran
O. isphaganica WC Iran, mountains around Esfahan city
O. laristanica Laristan and Fars province, Iran
Ovis vignei group = Urial
O. vignei, Ladakh, Kashmir in Pakistan and India and Wakhan and Ishkashim in Tajikistan
O. punjabiensis, Punjab province, Pakistan
O. bochariensis, NE Turkmenistan, S Uzbekistan, N Afghanistan, S Tajikistan
O. arabica, Oman and SE Saudi Arabia
O. cycloceros (including arkal) S Turkmenistan, NE Iran, Afghanistan
O. severtzovi Uzbekistan and the Turkestan range in Kyrgyzstan and Tajikistan
Ovis ammon group = Argali
O. nigrimontana Darya Karatau mountains, Kazakhstan
O. karelini, Tian-Shan mountains in SE Kazakhstan and E Kyrgyzstan, E to Urumqi in W China
O. collium Kazakhstan and NW China
O. polli E part of the Wakhan corridor, Afghanistan, Pamir plateau, Tajikistan, Pamir region and SE Kyrgyzstan into W China
O. hodgsoni, N India, N Nepal, N Bhutan, Tibet, NW Sichuan and SW Gansu and S Qinghai, China
O. ammon Altai mountains, W Mongolia, N Xinjang China, NE Kazakstan and Gorno-Alta and Tuvai republics in Siberian Russia
O. darwini S Mongolia, N China
O. jubata NE China
Sample sizes
Sample sizes for skins are not given
Skulls/horns (males only)
gmelini 3-7
isphaganica 1-8
laristanica 1-2
cycloceros 18-36
bochariensis 6-10
vignei 4-10
punjabiensis 1-6
arabica 0-1
severtzovi 0-5
nigrimontana 0-6
karelini 9-19
polli 8-31
collium 0-7
ammon 0-19
darwini 0-3
hodgsoni 2-16
The many 0 are due to the fact that the frontal arc was not measured in many specimens.
Data for jubata are taken from (Geist 1991) and are thus not shown in the main table.
Additionally there are samples from several hybrid populations (gmelini X arkal and cycloceros X laristanica)
Skins
The main difference between the vignei and gmelini group is that the gmelini group has a white saddle patch, which is absent in vignei. There are several minor pelage differences withing the gmelini group. The main difference between the vignei and ammon group is that the vignei group has a clear mane, absent in the ammon group.
Horns & skulls
vignei group
Both isphaganica and laristanica are said to have shorter teeth and thinner horns than gmelini, but there is wide overlap in teeth measurements and there is only 1 horn measurement for isphaganica and 2 for laristanica, so no reliable conclusion can be drawn
gmelini group
punjabiensis and bochariensis are very similar in skull characteristics and have on average a larger frontal arc than cycloceros, but there is overlap between bochariensis an cycloceros. The single arabica skull seems to have shorter teeth than all other taxa, the skull in general fits this group, though in terms of distribution a placement in the vignei group is more likely. in Skull heigth and skull breadth severtzovi seems somewhat intermediate between the gmelini and ammon group.
ammon group
ammon and hodgsoni are on average the largest taxa and this difference is absolute compared to some other taxa, though there are several intermediate taxa as well nigrimontana, darwini and collium are the smaller taxa. darwini is described as having a shorter, narrower but somewhat higher skull compared to hodgsoni and ammon, there is however large overlap in these measurements and sample sizes to small for reliable multivariate analyses. Overall a larger number of more subjective skull and horn measures are described, but given the data provided in the table combined with small sample sizes it is hard to draw generalizations from them.
Additional data
There has been quite some research on the genetics of wild sheep, especially because they are the forefathers of domestic sheep. There are clear differences genetically between the three groups. gmelini has a karyotype of 2n=54, vignei one of 2n=58 and ammon one of 2n=56. Research by Rezaei et al. (2010) indicate that ammon likely diverged from gmelini + vignei +- 1.72 million years ago (+-0.36 mya) and gmelini and vignei likely split 1.26 million years ago (+-0.36 mya). The relationships are however made more complex by extensive hybridization at the borders of these groups. severtzovi clearly groups in the ammon group. Several taxa: isphanicia, arkal, punjabiensis and gmelini are found not to be monophyletic based on genetic data.
Summarizing
Using a combination of morphological and genetic research it seems best to recognize three species of wild sheep in Eurasia next to the Snow sheep: Ovis gmelini (=Ovis orientalis) , Ovis vignei and Ovis ammon. Within each species there is quite some variation and all taxa elevated to species status by G&G can at least for now be recognized as subspecies, genetic research is however necessary to understand further the exact phylogenetic relationships within each group. Additionally the status of wild sheep on the Arabian peninsula is of conservation concern, as it is unclear whether they belong to vignei or gmelini.
Mouflon
Ovis gmelini gmelini
@Tomek , Tallinn Zoo, Estonia
Urial
Ovis vignei vignei
@Chlidonias , Hemis NP, India
Ovis vignei cycloceros (arkal)
@Nick@Amsterdam, Tierpark Berlin, Germany
Ovis vignei punjabiensis
@J I N X , Peshawar Zoo, Pakistan
Ovis vignei bochariensis
@Tomek , Zoo Liberec, Czechia
Argali
Ovis ammon ammon
@Giant Eland , Novosibirsk Zoo, Russia
Ovis ammon collium
@alexkant , Karaganda Zoo, Kazakhstan
Ovis ammon hodgsoni (including dalailamae)
@Deer Forest , Beijing Zoo
Ovis ammon jubata
@bongowwf , Shanghai Zoo, China
Ovis ammon polli
@Dormitator , Moscow Zoo breeding center, Sychovo, Russia
Ovis ammon darwini
@Chlidonias , Ikh Nart Nature Reserve, Mongolia
Hybrid sheep
Hybrid vignei X gmelini "Alborz sheep"
@fofo , Tehran Zoo, Iran
European mouflon
Ovis varies var. ophion Cyprus mouflon
@NigeW , Pafos Zoo, Cyprus
Ovis varies var. musimon European mouflon
@Tomek , New Forest Wildlife Park, UK
No pictures of nigrimontana, severtzovi, karelini, arabica, isphaganica or laristanica have been uploaded to the gallery yet
References
Hiendleder et al. (2002): https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1690972/pdf/12028771.pdf
Rezaei et al. (2010): https://www.sciencedirect.com/science/article/pii/S1055790309004461?via=ihubhttps://www.sciencedirect.com/science/article/pii/S1055790309004461?via=ihub
Sanna et al. (2016): The First Mitogenome of the Cyprus Mouflon (Ovis gmelini ophion): New Insights into the Phylogeny of the Genus Ovis
Next: the remaining Caprini
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What happened to the rest of the Capricornis species? Or Naemorhedus caudatus for that matter?
They will all be covered next, they are all part of the Caprini tribe, but they haven't been split by G&G.
Following Hassanin et al. (2012), the Caprini tribe covers all members of what used to be the Caprinae subfamily. Molecular data indicate that all Caprini are part of the Antilopinae subfamily, with Caprini being the sister to the Oryx and Hartebeest tribes.