Also added Arabian Tahr:
Arabian tahr (Arabitragus jayakari) - ZooChat
and possibly new subspecies of Salt's dik-dik and Kirk's dik-dik. I'd like your opinion:
Salt's dik-dik (Madoqua saltiana) - ZooChat
Kirk's dik-dik (Madoqua kirkii) - ZooChat
Ungulate taxonomy revisited: the evidence for the splits of G&G
- Thread starter lintworm
- Start date
Also added Arabian Tahr:
Arabian tahr (Arabitragus jayakari) - ZooChat
and possibly new subspecies of Salt's dik-dik and Kirk's dik-dik. I'd like your opinion:
Salt's dik-dik (Madoqua saltiana) - ZooChat
Kirk's dik-dik (Madoqua kirkii) - ZooChat
That paper is from December 2018. Below is a paper from June 2019 which appears to say the opposite (!), although I can only see the abstract.
ancient hybridization event reconciles mito-nuclear discordance among spiral-horned antelopes
The smallest of the spiral-horned antelopes, the bushbuck (T. scriptus), is also widely distributed across Africa, but is genetically divided into polyphyletic Scriptus and Sylvaticus mitochondrial (mt)DNA superlineages that inhabit opposite halves of the continent, suggesting the convergent evolution of independent bushbuck species. In this study, we provide a species tree reconstruction for the genus Tragelaphus and show that Scriptus and Sylvaticus are reciprocally monophyletic at nuclear DNA loci, comprising a single species across its African range. Given that mtDNA will sort into species-specific lineages more quickly than nuclear DNA, only an ancient interspecific hybridization event between a female from a now-extinct Tragelaphus species and a proto-Scriptus bushbuck male can reconcile the mito-nuclear incongruence.
Thanks for linking this study.
The interesting thing is that they come to a completely different conclusion with the same data. Hassanin et al. (2018) also found that they are monophyletic when it comes to nuclear dna, but not when it comes to mitochondrial dna. So the results are comparable. The conclusions drawn are not. This Rakotoarivelo et al. (2019) study argues that even though karyotypes differ, there is no stable hybrid zone, which is reflected by an earlier study and morphological data.
The conclusion drawn seems to depend mostly on the species concept used, where Rakotoarivelo et al. (2019) use a very strict Biological SC approach and Hassanin et al. (2018) a more practical approach. The Rakotoarivelo study simply ignores the long evolutionary separation (>1 million years) of the 2 main lineages as well as all the other clear differences described. Their caution is somewhat understandable because classifying Bushbuck in the contact area between scriptus and sylvaticus is difficult to almost impossible. I would however say that keeping 1 Bushbuck species is lumping a too large large amount of variation and evolutionary history in one taxa and only recognizing two separate species will make it generally clear what kind of variation is there, not only for taxonomists, but for everyone else who only looks at the species level.
Wapiti
Wapiti (or Elk for N-Americans) (Cervus canadensis) is a large species of deer from E Asia and N America and has in the past been treated as a subspecies of Red deer (Cervus elaphus). It is now widely regarded as a separate species and according to Mtdna the Sika deer (Cervus nippon s.l.) is the closest relative of the Wapiti.
There is a lot of discussion going on as to how many taxa exist within Wapiti and whether multiple species should be recognized, both in N America and Asia. IUCN recognizes the following taxa:
C.c. canadensis N America (including manitobensis, nelsoni)
C.c. nannodes California
C.c. roosevelti Vancouver Island, Washington and Oregon
C.c. alashanicus N China
C.c. sibiricus NE Kazakhstan, N Xinjang (China) to S Siberia and N Mongolia (includes songaricus)
C.c. xanthopygus SE Siberia, Russian Far East, Ussuriland & Manchuria,China
C.c. macneilli Central and SW China (including kansuensis)
C.c. wallichii SW China, Bhutan
In HMW Chinese Wapiti are split and macneilli and wallichi are treated as one separate species (including hanglu which has since been shown to be a separate species in the C. elaphus group).
G&G recognize 5 monotypic species
C. macneilli
C. wallichii
C. alashanicus
C. xanthopygus
C. canadensis including sibiricus, nannodes, roosevelti
Sample size
No sample sizes are given for antlers, skulls or skins, so it is impossible to verify data quality. Nor are quantitative anlalyses on skulls and antlers provided, so all we have are descriptions.
Skins
Some of the characteristics described by G&G include that both wallichii and macneilli are described as having a white rump patch, with the tail of wallichiii apparently somewhat shorter. Wallichii is yellowish-brown in winter, whereas macneilli is pale creamy grey with reddish tones. The rump patch of alashanicus is brownish white or russet orange, not overall coat colour is given. Rump patch of xanthopygus is described as reddish-yellow, shape and size resembling canadensis, which has a creamy rump patch. Stag xanthopygus are creamy-fawn in winter, canadensis is gray-brown or buff in winter. Some differences between canadensis from Asia are described, with animals from the Tianshan mountains (songaricus) and Altai (sibiricus) possibly differing in coat pattern and colour). No info is given on differences of canadensis within N America
Skulls & Antlers
No info is given at all about skulls or antlers of alashanicus. Both wallichii and macneilli appear to have a simpler antler than the other taxa, with wallichii having a 5 pointed antler and macneilli a 6 pointed antler. 3rd tine is disproportionally small in macneilli and less wide-spreading than wallichii. Both xanthopygus and canadensis have complex antlers, xanthopygus with 7 points, beam is somewhat curved in xanthopygus and curved in canadensis.
Additional data
A large body exists on the genetics of the Cervus genus, with increasing amounts of information.
One of the most recent papers by Meiri et al. (2018) shows that there are two main groups within Wapiti, a Chinese group consisting of xanthopygus, alashanicus, macneilli and wallichi and a second group consisting of sibiricus, songaricus and the N-American animals. This split is dated at about 0.3 million years ago (error margin: 0.14-0.47 mya). Meiri et al. (2018) find no evidence of genetic differences between songaricus and sibiricus, nor any structure within the N-American population. These results are the same as found by Kumar et al. (2016). Lorenzini & Garofalo(2015) find exactly the same pattern, but date the split between both groups at closer to 2 million years ago. I do not know enough about dating in such analyses to say anything informed about which one is more likely right or why they differ based on different methodologies. Intuitively I would however say that the real date of divergence is nearer to 0.3 than to 2 mya.
Meredith et al (2007) found genetic support for recognizing both nannodes and roosevelti as different from the nominate canadensis. Genetic research by Polziehn &Strobeck (2002); Polziehn et al. (1998) could not distinguish between canadensis and nelsoni, manitobensis, and merriami. But sibiricus can be identified as distinct from canadensis.
Summarizing
Based on the presented data there is no clear evidence to recognize the five proposed species, combined with DNA data there is however a case to be made whether one should recognize the Wapiti from China (wallichii, macneilli, alashanicus, xanthopygus) as a separate species (Shou). This group is genetically distinct from N-American wapiti + sibiricus, but it is unclear how distinct. Interestingly based on antler and colouration one would expect xanthopygus to be more closely related to canadensis, but it is apparently very small in size and this fits better with the other Shou (Chinese wapiti). Personally I think there is a pretty good case to be made to split Shou from Wapiti, but more data on nuclear dna and karyotype would be necessary to confirm such a split.
Cervus canadensis (canadensis?)
@Maguari , Tiergarten Nürnberg, Germany
@Maguari , Scottish Deer Centre, UK
Cervus canadensis sibiricus
@ThylacineAlive , Tierpark Berlin, Germany
@Maguari , Zoo Ostrava, Czechia
Cervus canadensis nannodes
@Dianamonkey , Zoo Plzen, Czechia
@Patrick87 , Tierpark Berlin, Germany
Cervus canadensis roosevelti
@Pleistohorse , Alaska Wildlife Conservation Center, USA
Cervus wallichii macneilli
@Deer Forest, Beijing Zoo, PR China
Cervus wallichii xanthopygus
@Deer Forest , Beijing Zoo, PR China
@Deer Forest , Badaling Wildlife Park, PR China
No pictures of alashanicus or wallichii have been uploaded to the gallery.
References
Kumar et al. (2016): https://www.researchgate.net/profil...g_the_phyl/links/5853887008ae7d33e01ac135.pdf
Lorenzini & Garofalo (2015): Error - Cookies Turned Off
Meiri et al. (2018): Error - Cookies Turned Off
Meredith et al. (2007): https://academic.oup.com/jmammal/article/88/3/801/1073029
Polziehn & Strobeck (2002): A Phylogenetic Comparison of Red Deer and Wapiti Using Mitochondrial DNA - ScienceDirect
Polziehn et al. (1998) http://www.friendsofkootenay.ca/sites/default/files/Polziehn et al 1998.pdf
Wapiti (or Elk for N-Americans) (Cervus canadensis) is a large species of deer from E Asia and N America and has in the past been treated as a subspecies of Red deer (Cervus elaphus). It is now widely regarded as a separate species and according to Mtdna the Sika deer (Cervus nippon s.l.) is the closest relative of the Wapiti.
There is a lot of discussion going on as to how many taxa exist within Wapiti and whether multiple species should be recognized, both in N America and Asia. IUCN recognizes the following taxa:
C.c. canadensis N America (including manitobensis, nelsoni)
C.c. nannodes California
C.c. roosevelti Vancouver Island, Washington and Oregon
C.c. alashanicus N China
C.c. sibiricus NE Kazakhstan, N Xinjang (China) to S Siberia and N Mongolia (includes songaricus)
C.c. xanthopygus SE Siberia, Russian Far East, Ussuriland & Manchuria,China
C.c. macneilli Central and SW China (including kansuensis)
C.c. wallichii SW China, Bhutan
In HMW Chinese Wapiti are split and macneilli and wallichi are treated as one separate species (including hanglu which has since been shown to be a separate species in the C. elaphus group).
G&G recognize 5 monotypic species
C. macneilli
C. wallichii
C. alashanicus
C. xanthopygus
C. canadensis including sibiricus, nannodes, roosevelti
Sample size
No sample sizes are given for antlers, skulls or skins, so it is impossible to verify data quality. Nor are quantitative anlalyses on skulls and antlers provided, so all we have are descriptions.
Skins
Some of the characteristics described by G&G include that both wallichii and macneilli are described as having a white rump patch, with the tail of wallichiii apparently somewhat shorter. Wallichii is yellowish-brown in winter, whereas macneilli is pale creamy grey with reddish tones. The rump patch of alashanicus is brownish white or russet orange, not overall coat colour is given. Rump patch of xanthopygus is described as reddish-yellow, shape and size resembling canadensis, which has a creamy rump patch. Stag xanthopygus are creamy-fawn in winter, canadensis is gray-brown or buff in winter. Some differences between canadensis from Asia are described, with animals from the Tianshan mountains (songaricus) and Altai (sibiricus) possibly differing in coat pattern and colour). No info is given on differences of canadensis within N America
Skulls & Antlers
No info is given at all about skulls or antlers of alashanicus. Both wallichii and macneilli appear to have a simpler antler than the other taxa, with wallichii having a 5 pointed antler and macneilli a 6 pointed antler. 3rd tine is disproportionally small in macneilli and less wide-spreading than wallichii. Both xanthopygus and canadensis have complex antlers, xanthopygus with 7 points, beam is somewhat curved in xanthopygus and curved in canadensis.
Additional data
A large body exists on the genetics of the Cervus genus, with increasing amounts of information.
One of the most recent papers by Meiri et al. (2018) shows that there are two main groups within Wapiti, a Chinese group consisting of xanthopygus, alashanicus, macneilli and wallichi and a second group consisting of sibiricus, songaricus and the N-American animals. This split is dated at about 0.3 million years ago (error margin: 0.14-0.47 mya). Meiri et al. (2018) find no evidence of genetic differences between songaricus and sibiricus, nor any structure within the N-American population. These results are the same as found by Kumar et al. (2016). Lorenzini & Garofalo(2015) find exactly the same pattern, but date the split between both groups at closer to 2 million years ago. I do not know enough about dating in such analyses to say anything informed about which one is more likely right or why they differ based on different methodologies. Intuitively I would however say that the real date of divergence is nearer to 0.3 than to 2 mya.
Meredith et al (2007) found genetic support for recognizing both nannodes and roosevelti as different from the nominate canadensis. Genetic research by Polziehn &Strobeck (2002); Polziehn et al. (1998) could not distinguish between canadensis and nelsoni, manitobensis, and merriami. But sibiricus can be identified as distinct from canadensis.
Summarizing
Based on the presented data there is no clear evidence to recognize the five proposed species, combined with DNA data there is however a case to be made whether one should recognize the Wapiti from China (wallichii, macneilli, alashanicus, xanthopygus) as a separate species (Shou). This group is genetically distinct from N-American wapiti + sibiricus, but it is unclear how distinct. Interestingly based on antler and colouration one would expect xanthopygus to be more closely related to canadensis, but it is apparently very small in size and this fits better with the other Shou (Chinese wapiti). Personally I think there is a pretty good case to be made to split Shou from Wapiti, but more data on nuclear dna and karyotype would be necessary to confirm such a split.
Cervus canadensis (canadensis?)
@Maguari , Tiergarten Nürnberg, Germany
@Maguari , Scottish Deer Centre, UK
Cervus canadensis sibiricus
@ThylacineAlive , Tierpark Berlin, Germany
@Maguari , Zoo Ostrava, Czechia
Cervus canadensis nannodes
@Dianamonkey , Zoo Plzen, Czechia
@Patrick87 , Tierpark Berlin, Germany
Cervus canadensis roosevelti
@Pleistohorse , Alaska Wildlife Conservation Center, USA
Cervus wallichii macneilli
@Deer Forest, Beijing Zoo, PR China
Cervus wallichii xanthopygus
@Deer Forest , Beijing Zoo, PR China
@Deer Forest , Badaling Wildlife Park, PR China
No pictures of alashanicus or wallichii have been uploaded to the gallery.
References
Kumar et al. (2016): https://www.researchgate.net/profil...g_the_phyl/links/5853887008ae7d33e01ac135.pdf
Lorenzini & Garofalo (2015): Error - Cookies Turned Off
Meiri et al. (2018): Error - Cookies Turned Off
Meredith et al. (2007): https://academic.oup.com/jmammal/article/88/3/801/1073029
Polziehn & Strobeck (2002): A Phylogenetic Comparison of Red Deer and Wapiti Using Mitochondrial DNA - ScienceDirect
Polziehn et al. (1998) http://www.friendsofkootenay.ca/sites/default/files/Polziehn et al 1998.pdf
DNA differences aside, none of these diverse Wapitis on the photos would be recognized as one, if put it in a large herd of Red Deer in any deer park in Europe.
Red Deer is a clear example how big is individual variability coupled with local feeding conditions. Which is completely missed if one looks at few skins or an inbred zoo line.
Wapiti
Wapiti (or Elk for N-Americans) (Cervus canadensis) is a large species of deer from E Asia and N America and has in the past been treated as a subspecies of Red deer (Cervus elaphus). It is now widely regarded as a separate species and according to Mtdna the Sika deer (Cervus nippon s.l.) is the closest relative of the Wapiti.
There is a lot of discussion going on as to how many taxa exist within Wapiti and whether multiple species should be recognized, both in N America and Asia. IUCN recognizes the following taxa:
C.c. canadensis N America (including manitobensis, nelsoni)
C.c. nannodes California
C.c. roosevelti Vancouver Island, Washington and Oregon
C.c. alashanicus N China
C.c. sibiricus NE Kazakhstan, N Xinjang (China) to S Siberia and N Mongolia (includes songaricus)
C.c. xanthopygus SE Siberia, Russian Far East, Ussuriland & Manchuria,China
C.c. macneilli Central and SW China (including kansuensis)
C.c. wallichii SW China, Bhutan
In HMW Chinese Wapiti are split and macneilli and wallichi are treated as one separate species (including hanglu which has since been shown to be a separate species in the C. elaphus group).
G&G recognize 5 monotypic species
C. macneilli
C. wallichii
C. alashanicus
C. xanthopygus
C. canadensis including sibiricus, nannodes, roosevelti
Sample size
No sample sizes are given for antlers, skulls or skins, so it is impossible to verify data quality. Nor are quantitative anlalyses on skulls and antlers provided, so all we have are descriptions.
Skins
Some of the characteristics described by G&G include that both wallichii and macneilli are described as having a white rump patch, with the tail of wallichiii apparently somewhat shorter. Wallichii is yellowish-brown in winter, whereas macneilli is pale creamy grey with reddish tones. The rump patch of alashanicus is brownish white or russet orange, not overall coat colour is given. Rump patch of xanthopygus is described as reddish-yellow, shape and size resembling canadensis, which has a creamy rump patch. Stag xanthopygus are creamy-fawn in winter, canadensis is gray-brown or buff in winter. Some differences between canadensis from Asia are described, with animals from the Tianshan mountains (songaricus) and Altai (sibiricus) possibly differing in coat pattern and colour). No info is given on differences of canadensis within N America
Skulls & Antlers
No info is given at all about skulls or antlers of alashanicus. Both wallichii and macneilli appear to have a simpler antler than the other taxa, with wallichii having a 5 pointed antler and macneilli a 6 pointed antler. 3rd tine is disproportionally small in macneilli and less wide-spreading than wallichii. Both xanthopygus and canadensis have complex antlers, xanthopygus with 7 points, beam is somewhat curved in xanthopygus and curved in canadensis.
Additional data
A large body exists on the genetics of the Cervus genus, with increasing amounts of information.
One of the most recent papers by Meiri et al. (2018) shows that there are two main groups within Wapiti, a Chinese group consisting of xanthopygus, alashanicus, macneilli and wallichi and a second group consisting of sibiricus, songaricus and the N-American animals. This split is dated at about 0.3 million years ago (error margin: 0.14-0.47 mya). Meiri et al. (2018) find no evidence of genetic differences between songaricus and sibiricus, nor any structure within the N-American population. These results are the same as found by Kumar et al. (2016). Lorenzini & Garofalo(2015) find exactly the same pattern, but date the split between both groups at closer to 2 million years ago. I do not know enough about dating in such analyses to say anything informed about which one is more likely right or why they differ based on different methodologies. Intuitively I would however say that the real date of divergence is nearer to 0.3 than to 2 mya.
Meredith et al (2007) found genetic support for recognizing both nannodes and roosevelti as different from the nominate canadensis. Genetic research by Polziehn &Strobeck (2002); Polziehn et al. (1998) could not distinguish between canadensis and nelsoni, manitobensis, and merriami. But sibiricus can be identified as distinct from canadensis.
Summarizing
Based on the presented data there is no clear evidence to recognize the five proposed species, combined with DNA data there is however a case to be made whether one should recognize the Wapiti from China (wallichii, macneilli, alashanicus, xanthopygus) as a separate species (Shou). This group is genetically distinct from N-American wapiti + sibiricus, but it is unclear how distinct. Interestingly based on antler and colouration one would expect xanthopygus to be more closely related to canadensis, but it is apparently very small in size and this fits better with the other Shou (Chinese wapiti). Personally I think there is a pretty good case to be made to split Shou from Wapiti, but more data on nuclear dna and karyotype would be necessary to confirm such a split.
Cervus canadensis (canadensis?)
@Maguari , Tiergarten Nürnberg, Germany
@Maguari , Scottish Deer Centre, UK
Cervus canadensis sibiricus
@ThylacineAlive , Tierpark Berlin, Germany
@Maguari , Zoo Ostrava, Czechia
Cervus canadensis nannodes
@Dianamonkey , Zoo Plzen, Czechia
@Patrick87 , Tierpark Berlin, Germany
Cervus canadensis roosevelti
@Pleistohorse , Alaska Wildlife Conservation Center, USA
Cervus wallichii macneilli
@Deer Forest, Beijing Zoo, PR China
Cervus wallichii xanthopygus
@Deer Forest , Beijing Zoo, PR China
@Deer Forest , Badaling Wildlife Park, PR China
No pictures of alashanicus or wallichii have been uploaded to the gallery.
References
Kumar et al. (2016): https://www.researchgate.net/profil...g_the_phyl/links/5853887008ae7d33e01ac135.pdf
Lorenzini & Garofalo (2015): Error - Cookies Turned Off
Meiri et al. (2018): Error - Cookies Turned Off
Meredith et al. (2007): Microsatellite Analysis of Three Subspecies of Elk ( Cervus elaphus ) in California
Polziehn & Strobeck (2002): A Phylogenetic Comparison of Red Deer and Wapiti Using Mitochondrial DNA - ScienceDirect
Polziehn et al. (1998) http://www.friendsofkootenay.ca/sites/default/files/Polziehn et al 1998.pdf
Another factor to consider with wapiti is that they have only been in North America for ~15,000 years (https://royalsocietypublishing.org/doi/full/10.1098/rspb.2013.2167), which makes it hard to justify recognition of multiple North American subspecies, let alone differentiation from sibiricus. As mentioned by @Jurek7 , a lot of the variation is probably the result of diet and climate. The Chinese forms certainly seem to differ enough both genetically and morphologically to represent a different entity, though.
You would only not recognize them if you were blind, Even the smallest Wapiti would be larger than the biggest Eurasian Red deer. Shou are somewhat smaller than Wapiti, but on average still clearly larger than Red deer. And that ignores obvious differences in the shape of the antlers and additionally differences in rutting calls have been described between different populations, which both point to a lot more divergence of populations than is to be expected by mere individiual differences based on local feeding situation. Variability in morphology within these populations is much smaller than between the populations. That is also to be expected if you take into account for how long these populations have developed separately. With the exception of the N-American Wapiti you would be the only one with such a view.
Also do you have any evidence for claiming all data comes from a few skins and inbred zoo lines? G&G seem to have seen mainly skins from the wild, when they state their sources....
Tule elk (nannodes) are genetically identifiable based on Meredith et al, (2007) and Polziehn & Strohbeck (2002), though they are not in the study by Meiri et al (2018), as well as morphologically. These analyses also show that American Wapiti can be genetically distinguished from sibiricus. nannodes is probably so unique genetically because of a genetic bottleneck. With small genetic differences as well as morphological differences between populations it would make sense to me to regard them as separate subspecies, as otherwise this variation is lost in the bigger picture. How much different populations have to differ to be regarded as separate subspecies or not is a matter of debate, but just blaming it on differences in diet is too easy.
Here's a photo I took in Mongolia of wild sibiricus - very different to European Red Deer at even a casual glance!
Sambar
Sambar (Cervus unicolor) are a species of deer widely distributed through S and E Asia. Sambar have been placed in the genus Rusa but genetic research has indicated that Rusa is nested within Cervus and is thus not a separate genus. A large number of subspecies have been described, but up to 7 are currently recognized:
C.u. unicolor Indian subcontinent, Sri Lanka, Bangladesh
C.u. dejeani C China
C.u. cambojensis From SE China through Indochina, Myanmar to Peninsular Malaysia
C.u. hainana Hainan Island, China
C.u. swinhoii, Taiwan
C.u. equina Sumatra
C.u. brookei Borneo
In HMW hainana and dejeani re not recognized.
G&G split Sambar in 2 species:
C. unicolor from Sri Lanka and the Indian subcontinent
C. equinus from SE Asia, Indonesia, China and Taiwan, as well as NE India (Assam & Bengal)
Sample sizes
G&G do not provide any sample sizes for skins, skulls or antlers, nor are any quantitative data presented.
Skins
Colour of unicolor is described as brown, varying from greyish to very dark brown in winter, more yellowish red brown in summer, belly darker than the back. Whereas equinus is described as dark brown in winter, more yellowish or reddish in summer. Tail is long-haired, very bushy in unicolor and even more bushy in equinus. Fawns of unicolor are spotted, whereas equinus lose their spotting rapidly with age.
Skulls & antlers
equinus is described as being somewhat smaller than unicolor, no skull characteristics of equinus are given, but unicolor is described as having a flat forehead and very deep preorbital fossa. In the antler the brow tine of unicolor is very short, compared to longer in equinus, as well as some other smaller differences in antler form.
Additional data
There have been surprisingly little studies on the differences in dna between different Sambar populations, even though there is a reason to expect some differences.
There seem to be quite some difference in karyotype (number of chromosomes) between different populations, with Leslie (2011) mentioning karyotypes of 2n = 56 for New Zealand animals (original from Sri Lanka & India), 2n=58 in India, 2n=62 for Malaysia and China. Additional studies give 2n=62 for Bornean animals (Idris & Moin 2009). G&G give 2n=60 as karyotype for their Indian species, but it is unclear to me where that comes from.
Summarizing
Overall there seems to be some variation between the different locations in karyotype, mainly between India and the rest, which corresponds with the two proposed species by G&G. But currently the “data” G&G provide are unsufficient as the basis for a split and extensive morphological, but more importantly genetic comparisons across the whole range of this species are highly needed. For now there is no evidence to back-up a split, but there are indications that this could change in the future.
References
Idris & Moin (2009): CAB Direct
Leslie (2011): https://watermark.silverchair.com/4...7x_yxLgCJ6VKyeKAG91--_ir0ptb1RNCJQdHK91Yy9GkM
Cervus unicolor unicolor
@ThylacineAlive , Bronx Zoo, USA
@Chlidonias , Sariska Tiger Reserve, India
Cervus unicolor cambojensis
@Maguari , Saigon Zoo, Vietnam
@LaughingDove , Khao Yai NP, Thailand
Cervus unicolor brookei
@Deer Forest , Lok Kawi Wildlife Park, Malaysia
Cervus unicolor swinhoii
@Al , Taipei Zoo, Taipei
Animals at TP Berlin have been labelled equina, but I am not sure whether that is correct. No pictures of hainana and possibly dejeani have been uploaded to the gallery.
Sambar (Cervus unicolor) are a species of deer widely distributed through S and E Asia. Sambar have been placed in the genus Rusa but genetic research has indicated that Rusa is nested within Cervus and is thus not a separate genus. A large number of subspecies have been described, but up to 7 are currently recognized:
C.u. unicolor Indian subcontinent, Sri Lanka, Bangladesh
C.u. dejeani C China
C.u. cambojensis From SE China through Indochina, Myanmar to Peninsular Malaysia
C.u. hainana Hainan Island, China
C.u. swinhoii, Taiwan
C.u. equina Sumatra
C.u. brookei Borneo
In HMW hainana and dejeani re not recognized.
G&G split Sambar in 2 species:
C. unicolor from Sri Lanka and the Indian subcontinent
C. equinus from SE Asia, Indonesia, China and Taiwan, as well as NE India (Assam & Bengal)
Sample sizes
G&G do not provide any sample sizes for skins, skulls or antlers, nor are any quantitative data presented.
Skins
Colour of unicolor is described as brown, varying from greyish to very dark brown in winter, more yellowish red brown in summer, belly darker than the back. Whereas equinus is described as dark brown in winter, more yellowish or reddish in summer. Tail is long-haired, very bushy in unicolor and even more bushy in equinus. Fawns of unicolor are spotted, whereas equinus lose their spotting rapidly with age.
Skulls & antlers
equinus is described as being somewhat smaller than unicolor, no skull characteristics of equinus are given, but unicolor is described as having a flat forehead and very deep preorbital fossa. In the antler the brow tine of unicolor is very short, compared to longer in equinus, as well as some other smaller differences in antler form.
Additional data
There have been surprisingly little studies on the differences in dna between different Sambar populations, even though there is a reason to expect some differences.
There seem to be quite some difference in karyotype (number of chromosomes) between different populations, with Leslie (2011) mentioning karyotypes of 2n = 56 for New Zealand animals (original from Sri Lanka & India), 2n=58 in India, 2n=62 for Malaysia and China. Additional studies give 2n=62 for Bornean animals (Idris & Moin 2009). G&G give 2n=60 as karyotype for their Indian species, but it is unclear to me where that comes from.
Summarizing
Overall there seems to be some variation between the different locations in karyotype, mainly between India and the rest, which corresponds with the two proposed species by G&G. But currently the “data” G&G provide are unsufficient as the basis for a split and extensive morphological, but more importantly genetic comparisons across the whole range of this species are highly needed. For now there is no evidence to back-up a split, but there are indications that this could change in the future.
References
Idris & Moin (2009): CAB Direct
Leslie (2011): https://watermark.silverchair.com/4...7x_yxLgCJ6VKyeKAG91--_ir0ptb1RNCJQdHK91Yy9GkM
Cervus unicolor unicolor
@ThylacineAlive , Bronx Zoo, USA
@Chlidonias , Sariska Tiger Reserve, India
Cervus unicolor cambojensis
@Maguari , Saigon Zoo, Vietnam
@LaughingDove , Khao Yai NP, Thailand
Cervus unicolor brookei
@Deer Forest , Lok Kawi Wildlife Park, Malaysia
Cervus unicolor swinhoii
@Al , Taipei Zoo, Taipei
Animals at TP Berlin have been labelled equina, but I am not sure whether that is correct. No pictures of hainana and possibly dejeani have been uploaded to the gallery.
Philippine deer
The Philippine deer (Cervus mariannus) has formerly been included in the genus Rusa, but based on genetic analyses, Rusa is nested within Cervus and should be regarded as a subgenus. Up to 4 subspecies have been recognized:
C.m. mariannus Luzon Island
C.m. nigricans lowlands of Mindanao Island and Basilan
C.m. nigella Upland Mindanao
C.m. barandanus Mindoro
G&G recognize three species, lumping nigricans with mariannus and elevating nigella and barandanus to species status.
Sample size
G&G base their split on an earlier paper by them from 1983, unfortunately this paper is not available online anywhere, so I cannot assess how many samples they had at their disposal. Based on a later paper (Meijaard & Groves 2004), they likely had no more than 14 male and 7 female samples at their disposal, which is very little.
Skins
No differences in skin between mariannus and barandanus are described. Pelage of nigella is described as soft, with a higher frequency of neck hair reversal than in lowland deer.
Skulls & antlers
Nigellus is described as very small. Barandanus is said to differ because of its unique width of braincase, mastoids, nasals and toothrow which are all “consistently and distinctively less” than in mariannus, as well as having a wider palate.
Additional data
No genetic research seems to have been done on this species.
Summarizing
This species is hardly known and should be a target species to understand the relationships between islands and within islands. Based on the evidence presented now by G&G it is unlikely we are dealing with multiple species, unless when seeing everything through an extremely narrow PSC look.
References
Grubb & Groves (1983): Notes on the taxonomy of the deer (Mammalia, Cervidae) of the Philippines
Meijaard & Groves (2004): ZSL Publications
Cervus mariannus (likely nominate)
@alexkant , Avilon Zoo, Philippines
@Giant Eland , Avilon Zoo, Philippines
The Philippine deer (Cervus mariannus) has formerly been included in the genus Rusa, but based on genetic analyses, Rusa is nested within Cervus and should be regarded as a subgenus. Up to 4 subspecies have been recognized:
C.m. mariannus Luzon Island
C.m. nigricans lowlands of Mindanao Island and Basilan
C.m. nigella Upland Mindanao
C.m. barandanus Mindoro
G&G recognize three species, lumping nigricans with mariannus and elevating nigella and barandanus to species status.
Sample size
G&G base their split on an earlier paper by them from 1983, unfortunately this paper is not available online anywhere, so I cannot assess how many samples they had at their disposal. Based on a later paper (Meijaard & Groves 2004), they likely had no more than 14 male and 7 female samples at their disposal, which is very little.
Skins
No differences in skin between mariannus and barandanus are described. Pelage of nigella is described as soft, with a higher frequency of neck hair reversal than in lowland deer.
Skulls & antlers
Nigellus is described as very small. Barandanus is said to differ because of its unique width of braincase, mastoids, nasals and toothrow which are all “consistently and distinctively less” than in mariannus, as well as having a wider palate.
Additional data
No genetic research seems to have been done on this species.
Summarizing
This species is hardly known and should be a target species to understand the relationships between islands and within islands. Based on the evidence presented now by G&G it is unlikely we are dealing with multiple species, unless when seeing everything through an extremely narrow PSC look.
References
Grubb & Groves (1983): Notes on the taxonomy of the deer (Mammalia, Cervidae) of the Philippines
Meijaard & Groves (2004): ZSL Publications
Cervus mariannus (likely nominate)
@alexkant , Avilon Zoo, Philippines
@Giant Eland , Avilon Zoo, Philippines
Now on to the remaining Cervini
Philippine spotted deer
Philippine spotted deer (Cervus alfredi) is the monotypic sister species to the Philippine (brown) deer (Cervus marianna). This species is restricted to Leyte, Cebu, Guimaras, Negros and neighboring islands in the Philippines.
@Bwassa, Edinburgh Zoo, UK
Rusa
The Rusa/ Javan deer / Timor deer (Cervus timorensis) is a small deer species that is native to Java and Bali. In the past millenia it has been introduced to many other Indonesian islands. Despite the fact that differences between islands have been described, Rusa are generally considered monotypic.
@Tomek , Lodz Zoo, Poland
@Giant Eland , Tamar Safari Bogor, Indonesia
White-lipped deer
The White-lipped or Thorold's deer (Cervus albirostris) is a large monotypic deer species from Central China (Tibet, Gansu, Xinghai, Yunnan). It has often been placed in the monotypic genus Przewalskium but genetic research indicates it is sister to the Red deer species group.
@ThylacineAlive , Highland Wildlife Park, Kingussie, UK
@Jogy , Tierpark Berlin, Germany
Pere David's deer
The Pere David's deer (Elaphurus davidianus) is a monotypic deer species with a former distribution on the coastal plains of N and C China, but is currently considered extinct in the wild, though re-introduction efforts are underway. The taxonomic placement of this species has long been difficult, but it is now accepted that it is the sister species to Eld's deer and these two species are the sister group of the genus Cervus.
@vogelcommando , Safaripark Beekse Bergen, Hilvarenbeek, the Netherlands
@nikola , Zoo Wroclaw, Poland
Philippine spotted deer
Philippine spotted deer (Cervus alfredi) is the monotypic sister species to the Philippine (brown) deer (Cervus marianna). This species is restricted to Leyte, Cebu, Guimaras, Negros and neighboring islands in the Philippines.
@Bwassa, Edinburgh Zoo, UK
Rusa
The Rusa/ Javan deer / Timor deer (Cervus timorensis) is a small deer species that is native to Java and Bali. In the past millenia it has been introduced to many other Indonesian islands. Despite the fact that differences between islands have been described, Rusa are generally considered monotypic.
@Tomek , Lodz Zoo, Poland
@Giant Eland , Tamar Safari Bogor, Indonesia
White-lipped deer
The White-lipped or Thorold's deer (Cervus albirostris) is a large monotypic deer species from Central China (Tibet, Gansu, Xinghai, Yunnan). It has often been placed in the monotypic genus Przewalskium but genetic research indicates it is sister to the Red deer species group.
@ThylacineAlive , Highland Wildlife Park, Kingussie, UK
@Jogy , Tierpark Berlin, Germany
Pere David's deer
The Pere David's deer (Elaphurus davidianus) is a monotypic deer species with a former distribution on the coastal plains of N and C China, but is currently considered extinct in the wild, though re-introduction efforts are underway. The taxonomic placement of this species has long been difficult, but it is now accepted that it is the sister species to Eld's deer and these two species are the sister group of the genus Cervus.
@vogelcommando , Safaripark Beekse Bergen, Hilvarenbeek, the Netherlands
@nikola , Zoo Wroclaw, Poland
And now the remaining Muntjac species.
Hairy-fronted muntjac
The Hairy-fronted muntjac or Black muntjac (Muntiacus crinifrons) is a monotypic species from E China.
@Giant Eland , Beijing Zoo, China
@Giant Eland , Beijing Zoo, China
Gongshan muntjac
The Gonshang muntjac (Muntiacus gongshanensis) is a recently described sister species of the Hairy-fronted muntjac. It occurs in S Yunnan, China and N Myanmar and possibly also in SE Tibet, NE India and Bhutan.
No pictures of this species have been uploaded to the gallery
Fea's muntjac
Fea's muntjac (Muntiacus feae) is a monotypic species from S Thailand.
@Giant Eland , Khao Kheow Forest & Wildlife Reserve Park, Thailand
@LaughingDove , Dusit Zoo, Thailand
Giant muntjac
The Giant or Large-antlered muntjac (Muntiacus vuquangensis) is a recently described species from Vietnam, E Laos and extreme E Cambodia.
No pictures of this species have been uploaded to the gallery.
Roosevelt's muntjac
Roosevelt's muntjac (Muntiacus rooseveltorum) is a small species from N Laos, but possibly also occurs in Vietnam and S China.
No pictures of this species have been uploaded to the gallery.
Leaf muntjac
The Leaf muntjac (Muntiacus putaoensis) is a small recently described species from N Myanmar and extreme NE India.
No pictures of this species have been uploaded to the gallery
Annamite muntjac
The Annamite muntjac (Muntiacus truongsonensis) is a small monotypic species that was recently described and occures in the S part of the Annamite range in Vietnam & Laos.
No pictures of this species have been uploaded to the gallery
Puhoat muntjac
The Puhoat muntjac (Muntiacus puhoatensis) is a recently described monotypic Muntjac species, known only from a tiny area in W Vietnam.
No pictures of this species have been uploaded to the gallery.
Reeves' muntjac
The Chinese or Reeves' muntjac (Muntiacus reevesi) is a small Muntjac species from SE China and Taiwan. Up to 6 subspecies have been proposed, of which 2 are widely recognized. The nominate species was introduced to Europe.
M.r. reevesi from mainland China
M.r. micrurus from Taiwan
Muntiacus reevesi reevesi
@Himimomi , Guangzhou Zoo, China
Muntiacus reevesi micrurus
@Maguari , Zoo Krefeld, Germany
Bornean yellow muntjac
The Bornean yellow muntjac (Muntiacus atherodes) is a monotypic species endemic to the island of Borneo.
No pictures of this species have been uploaded to the gallery.
Hairy-fronted muntjac
The Hairy-fronted muntjac or Black muntjac (Muntiacus crinifrons) is a monotypic species from E China.
@Giant Eland , Beijing Zoo, China
@Giant Eland , Beijing Zoo, China
Gongshan muntjac
The Gonshang muntjac (Muntiacus gongshanensis) is a recently described sister species of the Hairy-fronted muntjac. It occurs in S Yunnan, China and N Myanmar and possibly also in SE Tibet, NE India and Bhutan.
No pictures of this species have been uploaded to the gallery
Fea's muntjac
Fea's muntjac (Muntiacus feae) is a monotypic species from S Thailand.
@Giant Eland , Khao Kheow Forest & Wildlife Reserve Park, Thailand
@LaughingDove , Dusit Zoo, Thailand
Giant muntjac
The Giant or Large-antlered muntjac (Muntiacus vuquangensis) is a recently described species from Vietnam, E Laos and extreme E Cambodia.
No pictures of this species have been uploaded to the gallery.
Roosevelt's muntjac
Roosevelt's muntjac (Muntiacus rooseveltorum) is a small species from N Laos, but possibly also occurs in Vietnam and S China.
No pictures of this species have been uploaded to the gallery.
Leaf muntjac
The Leaf muntjac (Muntiacus putaoensis) is a small recently described species from N Myanmar and extreme NE India.
No pictures of this species have been uploaded to the gallery
Annamite muntjac
The Annamite muntjac (Muntiacus truongsonensis) is a small monotypic species that was recently described and occures in the S part of the Annamite range in Vietnam & Laos.
No pictures of this species have been uploaded to the gallery
Puhoat muntjac
The Puhoat muntjac (Muntiacus puhoatensis) is a recently described monotypic Muntjac species, known only from a tiny area in W Vietnam.
No pictures of this species have been uploaded to the gallery.
Reeves' muntjac
The Chinese or Reeves' muntjac (Muntiacus reevesi) is a small Muntjac species from SE China and Taiwan. Up to 6 subspecies have been proposed, of which 2 are widely recognized. The nominate species was introduced to Europe.
M.r. reevesi from mainland China
M.r. micrurus from Taiwan
Muntiacus reevesi reevesi
@Himimomi , Guangzhou Zoo, China
Muntiacus reevesi micrurus
@Maguari , Zoo Krefeld, Germany
Bornean yellow muntjac
The Bornean yellow muntjac (Muntiacus atherodes) is a monotypic species endemic to the island of Borneo.
No pictures of this species have been uploaded to the gallery.
I finally got to inserting the new photos of @Chlidonias and @Giant Eland. Additionally a quick look for the gallery gave new / improved pictures of several taxa, thanks to @Gondwana , @Jogy and myself.
For nominate C. canadensis, ZooAmerica has some. Considering the Manitoba ssp is no longer considered valid any photos of those should work as well. If nothing else, I do have photos of wild elk you could use but I've fallen really behind on uploading so haven't put them up yet...
According to ZTL, Berlin's old sambar (now at Howletts) came from San Diego, who have confirmed equina. One huge issue for the US population is that the two main collections breeding the species-- Bronx and SDZSP-- have two different (sub)species.
EDIT: San Diego has both (sub)species actually, but I believe equina is the population they're trying to grow for some reason.
~Thylo
According to ZTL, Berlin's old sambar (now at Howletts) came from San Diego, who have confirmed equina. One huge issue for the US population is that the two main collections breeding the species-- Bronx and SDZSP-- have two different (sub)species.
EDIT: San Diego has both (sub)species actually, but I believe equina is the population they're trying to grow for some reason.
~Thylo
I've got a (bad) puku photo and (ok) Namibian springbok photo as well, but it looks like there aren't any galleries for Namibia or Zambia.
Meiri et al (2018) did actually include a couple nannodes samples. They come out separately but mixed in with other wapitis (one closest to a roosevelti and the other closest to a canadensis), albiet with a limited genetic data set. Your point about their bottleneck-induced distinctiveness is well taken, though, and it certainly makes sense to manage them separately from other wapiti. As you said, whether that means calling them a separate subspecies really depends on how subspecies are defined, and if anything there's even less consensus in how to define subspecies than there is in how to define species.
There is actually one genetic study that incorporates C. mariannus (Systematic relationships of five newly sequenced cervid species). It is very limited, but does include two Luzon specimens representing the nominate subspecies and one from Leyte, which should be nigricans. Interestingly they differ as much from one another as they do from alfredi, which makes sense conceptually since these deer would all have still been on separate islands even during ice ages when sea levels were lower. Way too little data to make anything of it from a taxonomic standpoint though.
Thanks, are you sure the equina from San Diego came from Sumatra?
That would be welcome, you can request new galleries here:
New Gallery Request Thread 2019 [Open]
That is probably why many scientists now keep referring to Evolutionary Significant Units, so they don't burn their hands at taxonomy
I had seen that study, but didn't realize they had samples from different islands, and dismissed it as interesting, but not very useful for this discussion. I have been reading quite some papers on deer taxonomy recently and I recall one paper stated that alfredi and mariannus represent two seperate arrivals of deer to the Philippines. I am not sure which one it was, but that would imply that one could expect differences within mariannus to be smaller than with alfredi. I can try to dig that up again...
No idea where they came from, no. I can try and find out but I'm not sure I'll get anything.
~Thylo
CAPREOLINI
The tribe Capreolini consists of three species in two separate genera, the 2 species of Roe deer (Capreolus) and the Chinese water deer (Hydropotes inermis).
European roe deer
The European roe deer (Capreolus capreolus) is a medium-sized deer that occurs throughout most of Europe. HMW regards this species as monotypic, but based on Mtdna multiple subspecies have been described. Based on morphology up to 12 subspecies were described, but validity of most is doubtful.
C.c. capreolus N, W & C Europe
C.c. transsylvanicus E Europe
C.c. italicus parts of Italy
C.c. garganta (S?) Iberian Peninsula.
The exact distribution of the subspecies is unclear, as is the status of Roe deer from the Caucasus, and in Eastern Europe there are clear traces of ancient hybridization with the Siberian roe deer (C. pygargus)
Capreolus capreolus capreolus
@Daniel Sörensen , Gripsholms Castle deer park, Sweden
@Eagle , Zoo Dortmund, Germany
Winter pelage:
@Joker1706 , Zoo Dortmund, Germany
Capreolus capreolus transsylvanicus
@alexkant , Zoo Targoviste, Romania
@alexkant , Zoo Bucharest, Romania
Siberian roe deer
The Siberian roe deer (Capreolus pygargus) is closely related to the European roe deer, but differs in chromosome number and there are clear morphological differences, even though both species have been in contact recently. There has been debate as to whether multiple subspecies of Siberian roe deer could be recognized, with smaller studies finding no pattern, whereas larger more recent studies find clear differences between W and E populations corresponding to the previously described subspecies pygargus and tianschanicus (including bedfordi). It gets confusing with the status of animals in the Urals, Yakutia (NE Russia) and Jeju Island, South Korea. The animals from Jeju Island group apart from tianschanicus which occurs on the Korean Peninsula, and appears most closely related to animals in Yakutia. Jeju animals have been described as ochracea, but their taxonomic status is still unclear.
Conservatively we can recognize at least 2 subspecies, with the status of parts of the populations unclarified:
C.p. pygargus roughly west of Lake Baikal
C.p. tianschanicus Tian Shan Mountains, Mongolia, China, Russia E of Lake Baikal (includes bedfordi)
Status of animals on Jeju Islands, Jakutia & Urals unresolved.
Capreolus pygargys pygargus
@Giant Eland , Zoo Novosibirsk, Russia
@Giant Eland , Zoo Novosibirsk, Russia
Capreolus pygargys tianschanicus
@Giant Eland , Beijing Zoo, China
@Deer Forest , Beijing Zoo, China
@Deer Forest , Beijing Zoo, China
Chinese water-deer
The Chinese water deer (or just Water deer) (Hydropotes inermis) is a small species of deer originally from E China and the Korean Peninsula, but it has been introduced to England and France. Traditionally two subspecies have been recognized:
H.i. inermis E China, currently restricted to eastern Yangtze Basin.
H.i. argyropus W parts of the Korean Peninsula.
Recent genetic and morphological studies have however questioned the validity of these subspecies and Water deer might better be treated as a monotypic species. Groves did however ignore this research and said that “There is no evidence for more than one species, though this should be re-examined.”, without giving any indication why that would be necessary.
@ro6ca66 , Whipsnade Zoo, UK
@Tomek , Berlin Zoo, Germany
The tribe Capreolini consists of three species in two separate genera, the 2 species of Roe deer (Capreolus) and the Chinese water deer (Hydropotes inermis).
European roe deer
The European roe deer (Capreolus capreolus) is a medium-sized deer that occurs throughout most of Europe. HMW regards this species as monotypic, but based on Mtdna multiple subspecies have been described. Based on morphology up to 12 subspecies were described, but validity of most is doubtful.
C.c. capreolus N, W & C Europe
C.c. transsylvanicus E Europe
C.c. italicus parts of Italy
C.c. garganta (S?) Iberian Peninsula.
The exact distribution of the subspecies is unclear, as is the status of Roe deer from the Caucasus, and in Eastern Europe there are clear traces of ancient hybridization with the Siberian roe deer (C. pygargus)
Capreolus capreolus capreolus
@Daniel Sörensen , Gripsholms Castle deer park, Sweden
@Eagle , Zoo Dortmund, Germany
Winter pelage:
@Joker1706 , Zoo Dortmund, Germany
Capreolus capreolus transsylvanicus
@alexkant , Zoo Targoviste, Romania
@alexkant , Zoo Bucharest, Romania
Siberian roe deer
The Siberian roe deer (Capreolus pygargus) is closely related to the European roe deer, but differs in chromosome number and there are clear morphological differences, even though both species have been in contact recently. There has been debate as to whether multiple subspecies of Siberian roe deer could be recognized, with smaller studies finding no pattern, whereas larger more recent studies find clear differences between W and E populations corresponding to the previously described subspecies pygargus and tianschanicus (including bedfordi). It gets confusing with the status of animals in the Urals, Yakutia (NE Russia) and Jeju Island, South Korea. The animals from Jeju Island group apart from tianschanicus which occurs on the Korean Peninsula, and appears most closely related to animals in Yakutia. Jeju animals have been described as ochracea, but their taxonomic status is still unclear.
Conservatively we can recognize at least 2 subspecies, with the status of parts of the populations unclarified:
C.p. pygargus roughly west of Lake Baikal
C.p. tianschanicus Tian Shan Mountains, Mongolia, China, Russia E of Lake Baikal (includes bedfordi)
Status of animals on Jeju Islands, Jakutia & Urals unresolved.
Capreolus pygargys pygargus
@Giant Eland , Zoo Novosibirsk, Russia
@Giant Eland , Zoo Novosibirsk, Russia
Capreolus pygargys tianschanicus
@Giant Eland , Beijing Zoo, China
@Deer Forest , Beijing Zoo, China
@Deer Forest , Beijing Zoo, China
Chinese water-deer
The Chinese water deer (or just Water deer) (Hydropotes inermis) is a small species of deer originally from E China and the Korean Peninsula, but it has been introduced to England and France. Traditionally two subspecies have been recognized:
H.i. inermis E China, currently restricted to eastern Yangtze Basin.
H.i. argyropus W parts of the Korean Peninsula.
Recent genetic and morphological studies have however questioned the validity of these subspecies and Water deer might better be treated as a monotypic species. Groves did however ignore this research and said that “There is no evidence for more than one species, though this should be re-examined.”, without giving any indication why that would be necessary.
@ro6ca66 , Whipsnade Zoo, UK
@Tomek , Berlin Zoo, Germany
A Wapiti, whatever subspecies or sex, would stand out markedly in any herd of Red deer. Apart from size(and shape), they have very different colouration too. I am not surprised to hear the closest relatives of Wapiti are now thought to be Sika- the squealing calls of Sika in rut bear considerable similarity to the bugling of Wapiti. But neither are similar to the roaring call of Red deer at the rut.
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The Sambar are at Port Lympne these days, having moved from Howletts. I was under the impression that Howletts have had them a very long time though- almost from inception of the park, though I don't know their source. Do you have any other details of their arrival as having been from Berlin or when?