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

You are correct off-course I still haven't found a way to source it however

Btw. As I am going on holiday, there won't be any updates the coming month, but mid-January we will talk about Springbok here

 

Springbok

The Springbok (Antidorcas marsupialis) is a gazelle-like antelope from the (semi-)arid landscapes in Southwestern Africa.



Historically Springbok have been separated in three separate subspecies by some authors, but some authors including Kingdon's Mammals of Africa do not recognize any subspecies for springbok. The reason for this is that most of the variation is supposed to be on a cline with smaller animals in the south and larger animals in the north and this variation has been linked with environmental conditions and not so much with heredity. Non surprisingly G&G elevate the three putative subspecies to full species:

A. marsupialis S of the Orange river
A. hofmeyri N of the Orange river
A. angolensis Angola

Sample sizes

G&G do not present any original data in Ungulate Taxonomy and build completely on a 1981 paper by Groves (which in the original text is referred to as Groves 1981c, this should be Groves 1981a, Groves 1981c is about Dorcas gazelles...)

Skins
marsupialis 9
hofmeyri 18
angolensis 13

Skulls & horns (males/females)
marsupialis 7-73/13-43
hofmeyri 7-10/6-9
angolensis 10-11/3-4

It should be noted that the high numbers for marsupialis are only present for two out of six measurements: horn length and skull length. Overall sample sizes are rather high, except for the skulls and horns of hofmeyri and angolensis with the number of female measurements for angolensis being particularly low.

Skins

angolensis has wide face stripes, whereas they are thin in hofmeyri and very thin in marsupialis. Additionally there are some small differences in the colour of the flank stripe, the nose spot and the extent of the brown on the forehead. Groves in 1981 notes that in the dry centre of the range (hofmeyri) skins are more pale, whereas in moister northern and southern reaches of it's range skins are more strongly marked, corresponding to marsupialis and angolensis. Skins from Botswana (hofmeyri) are somewhat in between hofmeyri from Namibia and marsupialis from the Orange Free State, though still have more in common with the Namibian skins.

Skulls

Skulls from marsupialis are on average slightly smaller than those from hofmeyri and angolensis, both in males and females. There is however no difference between the latter two, except that the single male and female samples from the Kaokoveld, which is grouped into hofmeyri are larger than all other samples, but the skins look like typical Namibian hofmeyri.

Horns

The main difference between the three taxa is the difference in horn length between males and females, which is supposedly particularly large in marsupialis (only 60-70% of the male horn length) and relatively small in angolensis (87%), with typical hofmeyri values in between the other two taxa (77-85%). Average female horn length is however the same for the samples form Botswana, Swakopmund (both hofmeyri) and angolensis and given the averages and standard deviations there is also no difference in horn length in male angolensis and hofmeyri from Botswana, though a very limited number of samples from other hofmeyri localities has longer male horn lengths than for the other taxa. For measurements as horn span and horn base breadth there is just as much intrataxa as intertaxa variation. For horn antero-posterior length there seems to be a nice north-south cline with southern locality being smallest.

Additional data

There has been only one paper that has tried to find an answer to whether the differences between different Springbok populations are genetically driven or driven by environmental factors (van Aswegen et al. 2012) but they only compared hofmeyri with marsupialis. Their results were inconclusive however, though they show that there might be a link between genetics and phenotype.

Summarizing

Though there are some differences between different Springbok populations, the differences are rather weak and there is no clear evidence that these differences are caused by genetics and are thus heritable. For now I see no indication why the proposed splits by G&G are justified, but the preliminary recognition of three subspecies is ok based on a combination of skin and horn and skull characteristics, though it should be noted that especially within hofmeyri there is quite some variation as well.

Antidorcas marsupialis marsupialis

@Kudu21 , Shamwari Private Game Reserve, South Africa

Non-subspecific captive animal
@ThylacineAlive , Fort Worth Zoo, USA

References
Groves 1981: http://www.zobodat.at/pdf/Zeitschrift-Saeugetierkunde_46_0189-0197.pdf

van Aswegen et al. 2012: https://www.sciencedirect.com/science/article/pii/S161650471100156X

Next: the remaining Antilopini from Africa.

 

Now for the remaining African Antilopini:

Speke's gazelle
Speke's gazelle (Gazella spekei) is a gazelle endemic to Somalia and Somaliland and is a sister to Mountain gazelles (G. gazella) and the Dorcas gazelle (G. dorcas), it is monotypic and G&G do support that.


@Ituri , Reid Park Zoo, USA


@Kakapo , San Diego Zoo

Cuvier's gazelle
Cuvier's gazelle (Gazella cuvieri) is a monotypic gazelle that occurs only in the mountains in the Maghreb from the Western Sahara east to Tunisia. It is closely related to the Slender-horned gazelle (G. leptoceros).


@Giant Eland , Larry Johnson's place, USA


@Kakapo , San Diego Zoo, USA

Slender-horned gazelle
The slender-horned gazelle (Gazella leptoceros) occurs in the sand dunes of the Sahara from Algeria to Egypt. Originally Groves (1969) recognized two seperate subspecies, leptoceros in Egypt and loderi in Libya and Algeria. loderi is reported to be larger, but with slightly shorter horns (29.8 +- 1.6 cm vs. 33.7 +-2.8 cm, on a sample size of 6 for each taxon). G&G do not list any subspecies for Slender-horned gazelle in their Ungulate Taxonomy book though. An interesting note is that the Cuvier's gazelle from Tunisia are reported to be very close in dna to Slender-horned gazelle and the Cuvier's gazelle from more western areas not. There might be an interesting case in studying the genetics of these two closely related species in more detail, though both species have different habitat preferences.


@ThylacineAlive Bronx Zoo, USA


@ThylacineAlive , Bronx Zoo, USA

Red-fronted gazelle
The Red-fronted gazelle (Eudorcas rufifrons) occurs throughout the Sahel region from Senegal east to the Nile in Sudan. Currently three subspecies are recognized rufifrons from Senegal to Mali, laevipes Niger to the Nile and kanuri south of Lake Chad. G&G retain these three subspecies as marginally distinct.


@Maguari , Warsaw Zoo, Poland


@geomorph , San Diego Zoo Safari Park, USA

Heuglin's gazelle
The Heuglin's gazelle (Eudorcas tilonura) occurs in NE Sudan, NW Ethiopia and W Eritrea and has sometimes been treated as a subspecies of either Red-fronted gazelle or Thomson's gazelle but is currently treated as a monospecific species.

Mongalla gazelle
The Mongalla gazelle (Eudorcas albonotata) occurs in E Sudan and extreme Western Ethiopia. Just like the Heuglin's gazelle it has been treated as a subspecies of either the Red-fronted gazelle or Thomson's gazelle, but has now been accepted as being a separate monospecific species

Soemmerring's gazelle
The Soemmerring's gazelle (Nanger soemmerringi) is the sister species to the Grant's gazelle species complex. It occurs in Somalia, Somaliland, Ethiopian drylands, Eritrea and Sudan. Kingdon recognizes three subspecies butteri in S Ethiopia, berberana in Somalia and NE Ethiopia and soemmerringi in Eritrea and Sudan. G&G do not mention any of these subspecies and do not assess variation within this species, though they mention they expect it to be large. Apart from the three currently recognized subspecies there is most probably a fourth subspecies which occurs on the Dahlak islands of Eritrea and is noticeably smaller than the mainland taxa and with different skull characteristics (Chiozzi 2014)


@ThylacineAlive , San Diego Zoo, USA


@Giant Eland , Larry Johnson's Place, USA

Dama gazelle
The Dama gazelle (Nanger dama) is a more familiar gazelle to zoo enthusiasts as several of the subspecies are kept in zoos. Traditionally three subspecies are recognized: mhorr (Mhorr gazelle) in North Africa, dama from Senegal to Lake Chad and ruficollis (Red-necked gazelle) from Lake Chad to Sudan. G&G follow this arrangement. Senn et al. (2014) do however have a different view on the taxonomic arrangement of the Dama gazelle. They did not find any genetic support to recognize any subspecies. This goes against the well-described cline in coat pattern from the Mhorr gazelle in the NW and the Red-necked gazelle in the East. They argue however that as Dama gazelles had a continuous range up to 500 years ago and they are know to make long movements in search of water, this makes a strong case for high connectivity, which is reflected in their DNA work. They even argue that it would make sense to lump all subspecies for conservation purposes.

Nanger dama mhorr

@vogelcommando , Diergaarde Blijdorp, Netherlands

Nanger dama ruficollis


@Andrew_NZP , Pittsburgh Zoo, USA

Grant's Gazelle species group
The Grant's gazelle was originally treated as one species with four subspecies, but based on DNA evidence it is now clear that it should be treated as a species group with three species:
Bright's gazelle (Nanger notata)
Peter's gazelle (Nanger petersii)
Grant's gazelle (Nanger granti) with two subspecies: granti and robertsi, the latter being largely confined to the Serengeti ecosystem

Nanger notata

@lintworm , Abiatta-Shalla NP, Ethiopia


@lintworm , Abiatta-Shalla NP, Ethiopia

Nanger petersii

@Hix , Mkomazi NP, Tanzania

Nanger granti granti

@Jogy , Serengeti NP, Tanzania


Nanger granti robertsi

@lintworm , Lake Nakuru NP, Kenya


@lintworm , Ol Pejeta Conservancy, Kenya

Dibatag
The Dibatag (Ammodorcas clarkei) looks superficially similar to the Gerenuk, but is an unique antelope of Somalia and the Somali region in Ethiopia. Little is known about this species but no subspecies are recognized.

References

Chiozzi et al. 2014: Just another island dwarf? Phenotypic distinctiveness in the poorly known Soemmerring's Gazelle, Nanger soemmerringii (Cetartiodactyla: Bovidae), of Dahlak Kebir Island | Biological Journal of the Linnean Society | Oxford Academic

Senn et al. 2014: Splitting or Lumping? A Conservation Dilemma Exemplified by the Critically Endangered Dama Gazelle (Nanger dama)

Next is the Raphicerini tribe with the Sharpe's grysbok.

 

Sharpe's grysbok

The Sharpe's grysbok (Raphicerus sharpei) is a small antelope that occurs in Southeastern Africa, from Northern South Africa north to Central Tanzania. The Sharpe's grysbok is treated as monospecific by Kingdon, though some authors recognize two subspecies:

colonicus south of the Zambezi river
sharpei north of the Zambezi river.

Not very surprisingly G&G recognize both subspecies as distinct species and mention that a third species probably occurs in Central and Western Zambia.

Sample sizes

Skins
Sample size not mentioned

Skulls
colonicus (5-6)
sharpei (4)
Zambia (5)

Horns
colonicus (2)
sharpei (none)
Zambia (3)

Sample sizes are very small for all taxa

Skins

colonicus is said to be richer and darker brown than sharpei with lighter more reddish legs and the underparts more buff and a more distinct boundary between back and underparts. The skins of the Zambia taxa is not described at all.

Skulls and horns

colonicus is said to be larger, but with a relatively more narrow skull. Though on average this difference is there, as shown in the measurements of gt1 and Cb1, the extremes of each subspecies overlap. There are no other differences in skull measurements. The putative Zambia taxa is said to be somewhat larger with smaller teeth, though this is not at all visible in the data they present. The only possible difference is is the nasal breadth which is somewhat larger, though the smallest Zambian measurement is the same as the maximum measurement for both colonicus and sharpei. There is no difference in horn length between the taxa.

Additional data

There does not seem to be any genetic data about possible differences between the Sharpe's grysbok taxa.

Summarizing

Based on the very small differences combined with the small sample sizes there seems to be no reason to recognize more than one species of Sharpe's grysbok. The recognition of subspecies seems justified as there are some minor pelage differences and based on the small sample there is some minimal difference in skull size and breadth, but none of these differences is absolute even with this small sample size.

No pictures have yet been uploaded to the gallery

Next the remaining Raphicerini: Cape grysbok and Steenbok.

 

Cape grysbok

The Cape grysbok (Raphicerus melanotis) is the sister species of the Sharpe's grysbok and is endemic to southern South Africa. This species has sometimes been lumped with the Sharpe's grysbok, but the Cape grysbok has lateral hooves, which are missing in Sharpe's grysbok, it also has longer horns and there are some differences in pelage.


@snowleopard , International Wildlife Museum, USA

Steenbok
The steenbok (Raphicerus campestris) is the most well known species in this genus. Though steenbok is Dutch for ibex, it looks nothing like an ibex... Steenbok have a disjunct distribution, occurring in Kenya, Tanzania & Uganda in the East and from Angola, Zambia and Mozambique south to the Cape. The species does not occur in the Miombo woodlands separating these two regions. Up to nine subspecies have been recognized, but Kingdon argues that it the most sensible solution is to recognize only two subspecies:

campestris in Southern Africa
neumanni in East Africa

G&G do not split the Steenbok in separate species, but do split the campestris subspecies in 5 subspecies:
campestris S and SW Cape province, South Africa
fulvorubescens Albany district, South Africa
natalensis (including zuluensis) Kwazulu Natal into the Transvaal, South Africa
capricornis between the Limpopo and Zambezi river, so South Africa north to Zambia
steinhardti (including kelleni) NW South Africa, Namibia, Botswana into Angola

Sample sizes

Skins
No sample sizes are reported.

Skulls
campestris (1-2)
fulvorubescens (1)
natalensis (11)
capricornis (6)
steinhardti (17-18)
neumanni (11)

Horns
campestris (1)
fulvorubescens (1)
natalensis (4)
capricornis (4)
steinhardti (7)
neumanni (9)

Sample sizes for horns are very small for most taxa and also skull sample sizes are extremely small for two proposed subspecies

Skins

G&G already note that the differences in external features are slight. natalensis and capricornis are "much the same" when it comes to the pelage. They are said to be more rufous and with a lighter face when compared to fulvorubescens with the white extending from the throat to the chin. fulvorubescens is said to be "near amber-brown". campestris is said to be more orange-rufous in general with cheeks and legs ochraceous-tawny, the white on the underparts is not extending down the inside of the limbs as in fulvorubescens and the white on the throat is also less extensive. steinhardti is again much the same as capricornis and natalensis but paler than the other taxa. neumanni from E Africa has larger white eye-rings and more white on the margins of the ears, the white is also more clearly defined on the underside as well as on the lips

Skulls & horns

There are no differences at all in the skull measurements, though there are some very minor differences on average, with campestris (one specimen only) having small nasal bones on average, but it falls well within the range of natalensis. natalensis is said to differ from capricornis by its larger horn size, but this is based on a very small sample size (though it is distinct between these two taxa, both fall within the range of other S African taxa.). There are no differences between the E African and Southern African populations.

Additional data

There does not seem to be any data on intraspecific genetics of Steenbok.

Summarizing

G&G recognize six instead of two subspecies of Steenbok. The evidence for this subspecific division is rather weak though and there does not be much support to recognize more than two subspecies. Even the recognition of two subspecies is debatable as the differences in pelage are rather small, even though there is a clear geographical barrier. Genetics could resolve whether the eastern population is indeed distinct from the southern population. For now I would keep the two subspecies neumanni and campestris provisionally, but more evidence is needed to see whether they are distinct enough to warrant recognition.

Raphicerus campestris campestris


@GerbenElzinga , Kruger NP, South Africa

Raphicerus campestris neumanni


@Hix , Serengeti NP, Tanzania

Next the Madoquini, Dikdiks, with Salt's dikdik coming first.

 

Salt's dik-dik

The Salt's dik-dik (Madoqua saltiana) is a dikdik that occurs in the Horn of Africa with the core of its distribution in Somalia, Somaliland and Ethiopia. Historically the Silver dik-dik (Madoqua piacentinii) has sometimes been treated as a subspecies of the Salt's dik-dik, but it now generally accepted as being a distinct species.

Generally five subspecies have been recognized, though Kingdon has noted that these might be actual species:

M.s. saltiana NE Sudan, Eritrea, Djibouti, Afar triangle (Ethiopia)
M.s. phillipsi N Somalia (Somaliland)
M.s. hararensis N Somalia (Somaliland), south of mountain range, Ogaden region Ethiopia to the Shebelle river
M.s. lawrancei coastal Somalia
M.s. swaynei S Ethiopia, S Somalia, south of the Shebelle river

Naturally G&G elevate all five subspecies to species status.

G&G do not seem to have done any extra research themselves and seem to base their conclusions solely on Yalden (1978), who did the last revision of the Salt's + Silver dik-dik.

Sample sizes

Skins
Sample sizes for skins are not reported

Skulls
saltiana 31
phillipsi 38
hararensis 12
lawrancei 22
swaynei 10

So sample sizes are quite good, but Yalden only checked one variable, the length of the upper tooth row.

Skins

saltiana is agouti-reddish gray coloured on the upperparts, with legs sandy coloured. philipsi is grayer than saltiana with the pale orange of the forelegs extending into flares over the shoulder. hararensis is said to be more reddish than saltiana and philipsi with the red spreading to the back. swaynei is coloured like saltiana but lawrencei is more silver-gray dorsally, sharply contrasting the deep reddish-brown legs, shoulders and flanks.

Skulls

Yalden statistically tested for the difference in the length of the upper tooth row, though I don't see which test he used. It is however clear that saltiana has a significantly longer tooth row than the other taxa, though there is overlap in the standard deviation with swaynei. phillipsi, hararensis and lawrancei have the same upper tooth row length. swaynei has a slightly larger average upper tooth row length than the aforementioned three taxa, but these differences are rather small and seem not significant.

Additional data

I haven't been able to find any DNA data on Salt's dikdik or other morphological analysis.

Summarizing

There is some clinal variation within the taxa (e.g. in phillipsi individuals from the east are more clearly marked than in the west), but very clearly between the taxa (also see Figure 5 in Yalden (1978)). There are some differences in the upper tooth length between the taxa (on which G&G base their "size" characteristics), but only the nominate saltiana seems to stick out. There are some additional differences in the coat pattern and with the Shebelle river and some highland areas there are potential barriers between most of the taxa (except hararensis and phillipsi). As we will see in the Kirk's dik-dik, there can be a lot more variation than is at first apparent in the Madoqua genus. The pelage differences and the slight differences on the one skull character measured, might indicate potential speciation. For this we however need data on their DNA and chromosome count and it would also be necessary to measure more skull characteristics. So for now it seems best to keep the Salt's dik-dik as one species, with five subspecies. In light of new evidence this could however change...

next: Guenther's dik-dik

Madoqua saltiana saltiana

@Maguari , Awash NP, Ethiopia


@lintworm , Awash NP, Ethiopia

Possibly Madoqua saltiana swaynei

@Giant Eland , Albustan Zoological Centre, UAE

Madoqua piacentinii

@alexkant , Moscow Zoo, Russia

No picture of the other Salt's dik-dik taxa have been uploaded.


Reference

Yalden 1978: https://www.tandfonline.com/doi/pdf/10.1080/03749444.1978.10736583

 

I have started to link to zoochat pictures of the various antelope taxa, to also have a visual reference. For now I only added pictures for Greater and Lesser kudu, but more will follow soon.

So if anyone has pictures from obscure African antelope taxa, preferably from the wild, please upload them to the gallery.

 

Günther's dik-dik

Günther's dik-dik (Madoqua guentheri) is the sister species to the Kirk's dikdik (Madoqua kirkii s.l.) species complex. M. guentheri and M. kirkii s.l. have hybridized in the past, but this did not result in fertile offspring. M. guentheri occurs in the driest areas of all Dikdik from NE Uganda, throughout N Kenya, S Ethiopia, Somalia to Somaliland. Up to five subspecies have been recognized, of which the validity of two is very doubtful. Two subspecies are generally recognized:

M.g. guentheri (including hodsoni, wroughtoni) Somalia, Somaliland SE Ethiopia, NE Kenya
M.g. smithii (including nasoguttatus) Uganda, N Kenya, S Ethiopia

note that Kingswood & Kumamoto (1996) give the Wabe Shebelle river (Somalia) as border between the two subspecies. This would extend the range of smithii significantly.

G&G elevate these two subspecies to species status. Interestingly HMW does not acknowledge this split.

Sample sizes

No sample sizes are given for skins and measurements on skull characteristics are taken from Kingswood & Kumamoto (1996):

guentheri 1-3
wroughtoni 1-6
nasoguttatus 1
smithii 1-13

it is not clear which sample size was present for each measurement

Skins
smithii is described as more reddish than guentheri and smithii is said to have buffy white underparts, which are pinkish buff in guentheri.

Skulls
The given average in Ungulate Taxonomy for greatest skull length is different than in Kingswood & Kumamoto. The measurements given by G&G are 98-109 mm for guentheri and 109-120 mm for smithii. Kingswood & Kumamoto however give ranges of 100-111 for guentheri and 110-120 for smithii. G&G thus falsely claim that the distinction in greatest skull length is absolute between guentheri and smithii. Overall smithii is larger than guentherii, this difference is absolute for condylobasal skull length 105.7 mm (100-112) vs. 95.2 mm (93-97), other differences are not absolute or sample size for either guentheri (including wroughtoni) or smithii (including nasoguttatus) is only 1, but on average smithii is always larger.

Additional data
I am not aware of any genetic data looking at the differences between smithii and guentherii. Interestingly the chromosome number can differ within a population of Günther's dikdik(either n=48 or n=50), but crossbreeding results in fertile offspring.

Summarizing
smithii and guentherii are not as distinct as G&G do claim in their book. On average smithii is still larger, but this difference was only absolute for one measurement and limited by a small sample size. Differences in pelage are rather limited, but differences between the different species in the Kirk's dikdik complex can also be small. It would be interesting to see genetic data across the range of this species to see whether there is any case to be made to split this species into two. For now recognizing one species with two subspecies smithii and guentheri seems the best option.

Reference:
Kingswood & Kumamoto (1996): https://watermark.silverchair.com/5...Yfy6FXx5vQBymq-L3hADYmRDWaVMYiGpe35oCcfoPrZCA

Madoqua guentheri smithii

@keith8404 , Tulsa Zoo, USA


@Dibatag , Calgary Zoo, Canada

Based on the limited amount of pictures the Günther's dik-dik in N America seem to belong to the smithii subspecies, but any information on the exact origin is welcome. smithii would also be more logical as it occurs in Kenya & Uganda, which have been a source of many antelopes in zoos, more so than Ethiopia and Somalia. So far no pictures of M.g. guentheri have been uploaded.

next: Kirk's dikdik

 

The population was of smithii, though the population is now down to a single animal in Texas.

~Thylo

 

Kirk's dik-dik

The Kirk's dik-dik (Madoqua kirkii) has traditionally been treated as one species with a disjunct distribution, occuring in East Africa and Namibia, Angola. In the 1990s it became clear that the Namibia population has a different chromosome count than part of the East African population and it was also found that there are also three different types of Kirk's dik-dik in East Africa, each with their own chromosome count. First hints that this could be the case came from San Diego where Kirk's dik-dik from two different localities gave only offspring where the males were sterile. It was later found out that these were actually two species: the Kirk's dik-dik and the Naivasha dik-dik (M. cavendishi).

Based on the genetic work the Kirk's dik-dik species complex has now four recognized species:

Madoqua kirkii (including minor, nyikae, 2n=47) NE African coast, from S Somalia to NE Tanzania, including most of E Kenya

Madoqua cavendishi (including hindei, 2n=46). E Uganda to C Kenya including N Tanzania

Madoqua thomasi (2n=48) Tanzania, except extreme N and NE

Madoqua damarensis (2n=48) NW Namibia and SW Angola

These four species are also recognized by G&G, they however also recognize a 5th species Madoqua hindei. This species is said to occur in the range described for M. kirkii south of the Tana river, thus M. kirkii being limited to NE Kenya and SW Somalia N of the Tana river. Confusingly hindei is described by Kingdon to be part of cavendishi. The type specimen for hindei comes from Kitui which lies more or less on the border between the known kirkii and cavendishi distributions. But Kingswood & Kumamoto (1997) describe hindei as a highland race, which kirkii is certainly not. It seems to me more likely that G&G should have named their 5th species nyikae, which was described from Voi, Kenya, which is part of the range G&G describe for hindei.

As four species of Kirk's dik-dik are well defined, the only real controversy is whether Madoqua (kirkii) kirkii should be split into a northern kirkii and a southern nyikae. Because it is somewhat confusing how the 5th species should be called, I will compare the measurements for nyikae, cavendishi, hindei and kirkii.

Sample sizes

Sample sizes for skins are not reported and no sample sizes are reported for skulls by G&G. I assume however that they based their analysis on the data published by Kingswood & Kumamoto (1997), plus additional skulls obtained from them for cavendishi and kirkii. Sample sizes reported here are purely based on Kingswood & Kumamoto (1997):

Skulls (males/females)
kirkii (4-5/0)
cavendishi (6-8/5-6)
hindei (2-3/1)
nyikae (6-8/2)

It is clear that sample sizes are really small, especially for females.

Skins

hindei (with nyikae as synonym) is described in a lot of detail by G&G but the main difference is that overall the skin colour is described as red-olive, compared to the paler kirkii and darker cavendishi. hindei also has less white on the underside, limbs and face.

Skulls

hindei is described by G&G as the only species where the males are larger than the females, but this is based on a single female measurement (109 mm), vs 3 from males (110-116 mm) of greatest skull length. nyikae samples show males (111.7 mm; 106-116) and females (113.5 mm; 112-115) to be almost similar. The kirkii males are all smaller with a maximum length of 106 mm and cavendishi are bigger with 116-127 mm. The second measurement: maxillary tooth row length shows however on average longer lengths for males compared to females for all taxa, though females are generally a bit larger in other measurements. Here hindei and nyikae males have the same length, though the single nyikae measurement for females is much smaller than the 2 hindei samples. kirkii males have on average smaller measurements, but the biggest length of kirkii (37 mm), falls within the range of both nyikae (34.4 mm as smallest measurement) and hindei (36.2 mm as smallest measurement). cavendishi is on average slightly larger, but not on absolute values.

Additional data

I am not aware of any additional data, apart from the already mentioned dna data that looked at chromosome numbers.

Summarizing

It is a bit of a mess here, because there is no DNA evidence that a fifth species of Kirk's dik-dik is valid. There are some differences in pelage for hindei and in terms of skull size hindei/nyikae falls in between kirkii and cavendishi. The claim that hindei males are larger than their females, is however based on nothing. Overall G&G have not been able to make a strong case as to why hindei is different from kirkii especially given the lack of DNA support and the extremely small sample sizes. So personally the status of recognizing 4 species of Kirk's dik-dik, potentially with subspecies seems fair enough.

Madoqua kirkii:


@lintworm , Samburu National Reserve, Kenya



@ro6ca66 Cotswold Wildlife Park, UK

Madoqua kirkii nyikae ("Madoqua hindei" for G&G)

@lintworm , Mkomazi NP, Tanzania

Madoqua cavendishi


@geomorph , San Diego Safari Park, USA


@lintworm , Lake Nakuru NP, Kenya

Madoqua thomasi

@lintworm , Ruaha NP, Tanzania


@LaughingDove , Tarangire NP, Tanzania

No pictures available for Madoqua damarensis

Reference:

Kingswood & Kumamoto (1997): https://watermark.silverchair.com/5...2NmdaoGg0LW4YtSqLpfM1swKcuMLm-3xxW8VUI19Vf2_g

The next antelope will be the one we have all been waiting for:
Klipspringer

 

Klipspringer

The Klipspringer (Oreotragus oreotragus) is a unique antelope that is confined to rocky outcrops and mountain areas throughout Southern and Eastern Africa, with a relict population in Nigeria and the CAR. Due to its habitat requirements it has a rather patchy distribution and up to 11 subspecies have generally been recognized, though their validity has been questioned:

O. o. oreotragus Cape region and E Free State, South Africa
O. o. transvaalensis N and E South Africa
O. o. tyleri Namibia and SW Angola
O. o. stevensoni Zimbabwe and E Botswana
O. o. aceratos SE Africa between the Rufiji and Zambezi rivers
O. o. centralis Zambia, DR Congo, SW Tanzania, NW Malawi
O. o. schillingsi most of Tanzania, S Kenya, SW Uganda
O. o. aureus N Kenya, NE Uganda, S Ethiopia
O. o. saltatraxoides Ethiopian highlands, Eritrea, E Sudan
O. o. somalicus N Somalia and Djibouti
O. o. porteousi Nigeria and Central African Republic

All these subspecies are elevated to species level by G&G but the species range listed is much smaller for most taxa in Ungulate Taxonomy, correct distributions are given by Castello, but the wrong distributions are given in HMW, implying clear boundaries between the taxa and thus ignoring areas where Klipspringer do occur (e.g. Northern Ethiopian highlands, most of Namibia etc.).

Sample sizes

No sample sizes are given for skins, for skulls and horns the following sample sizes are given (note that only in schillingsi females generally have horns)

Skulls and horns (males/females)
porteousi 2-4 / 0
Sudan 1 / 2
somalicus 0 / 3
saltatrixoides 7-8 / 3
Ankole 2 / 3
NW Kenya 4-5 / 1
aureus 7 / 7
Kenyan highlands 4-19 / 1
schillingsi 4-6 / 3-4
aceratos 1 / 3
centralis 16 / 7
W Zambia 1 / 1
stevensoni 4-5 / 1
tyleri 8-11 / 8-9
transvaalensis 3 / 2
oreotragus 2-3 / 3-4

Note that Kenyan highlands, NW Kenya and Ankole are lumped with aureus, Sudan is lumped with saltatraxoides and W Zambia is lumped with centralis

Overall sample sizes are small to very small for most taxa. Only centralis, aureus and tyleri are moderately to well represented.

Skins
oreotragus is described as uniform yellow with a pale underside. Neighbouring tyleri is pale sandy-ochre, with underparts broadly uniformly white. transvaalensis is relatively similar but more brightly yellow and with whiter underparts than oreotragus. stevensoni is duller in colour than transvaalensis and darker, especially the head. centralis rufous to yellow-gray and lacks the black/dark present above the forehoofs which is present in the southern taxa. aceratos has ochery forequarters and olive hindquarters. massaicus and saltatraxoides are very similar with a golden orange color. aureus is more golden-yellow and has a rufous crown. somalicus is yellowy-olive with a brown crown and white underparts. porteousi is variable in colour but more dull yellowy. For porteousi, saltatrixoides and schillingsi the variation within each taxa is more variable than between the taxa.

Horns and skulls
schillingsi is the only taxa in which the females also have horns. Many other differences are also described by G&G between the taxa, but these differences are a) based on very small sample sizes , b) only hold up when comparing averages, without taking into account the range of values within each taxa and c) taxa with high number of samples show a large variation. It thus seems reasonable to assume that if more samples become available for other taxa, their range in measurements will also increase. There are a few things that become apparent though: the females in oreotragus are relatively small, especially when compared to the size of oreotragus males. transvaalensis have relatively large horns and the animals from Sudan are very small (but only 3 samples available). for schillingsi females not only have horns, but they are also larger. G&G further describe a lot of differences in size and teeth size, but the conclusions they draw, cannot possibly be drawn from the available data.

Additional data
Robinson et al. (1995) describe that there is no difference in cytotype between different Southern African taxa (probably oreotragus and transvaalensis), but they also note that this doesn't mean there is no genetic difference between the taxa. For her MSc thesis Le Roex sampled DNA of 9 of the 11 taxa (missing only porteousi and stevensoni). She showed a split between southern and eastern populations and a split within the Southern populations. The Southern versus Eastern divergence is as high as 11%, comparable with e.g. Grant's gazelle (which have now been split) and Kob (not split). The two Southern groups differ about 6%. Most subspecies could also be recognized based on their dna, only transvaalensis and aceratos were not distinguishable, so could be merged.

Summarizing
Based on the work by G&G there is no compelling reason to split Klipspringers, their conclusions are drawn from a very small sample size. The work by Le Roex is however very interesting and raises the possibility that there are at least two Klipspringer species, one in Southern Africa and the other in Eastern Africa. The position of the Nigerian and Central African Populations remains an enigma though. Le Roex's work was however based on small sample sizes for many subspecies, so additional work is necessary to see whether the subspecific differences would be justified in all cases. It is also of high conservation importance to include porteousi in that analysis, as it is the most threatened taxa and possibly a separate species. The evidence seems strong enough to recognize two Klipspringer species within this species group (with the possibility of a third in Nigeria). So then we would provisionally get:

Oreotragus (o.) oreotragus (Zimmermann 1783) with the following subspecies:
oreotragus
stevensoni
centralis
tyleri
aceratos

Oreotragus (o.) saltatrixoides (Temminck. 1853) with the following subspecies
saltatrixoides
schillingsi
somalicus
aureus
porteousi

Oreotragus oreotragus oreotragus

@Gondwana , Western Cape, South Africa

Oreotragus oreotragus aceratos (formerly transvaalensis)

@LaughingDove , Entabeni Game Reserve, South Africa



Oreotragus saltatrixoides saltatrixoides

@MagpieGoose , Biopark Valencia, Spain


@Javan Rhino , Zoo Frankfurt, Germany


Oreotragus saltatrixoided schillingsii

@Hix , Tarangire NP, Tanzania


References

Le Roex (2008) http://open.uct.ac.za/bitstream/handle/11427/19029/thesis_sci_2008_le_roex_nikki.pdf?sequence=1

Robinson et al. (1995) http://www.zobodat.at/pdf/Zeitschrift-Saeugetierkunde_61_0049-0053.pdf

Next: Suni

 

So by now I have added photos for all taxa for which photos were available in the gallery. I should have some pictures of hofmeyri springbok, Northern gerenuk and Damara dik-dik. If anybody sees something he/she has, please upload. So for now I already want to thank @molinea , @sebbe67 , @Maguari , @Tim May , @Tomek , @jbnbsn99 , @ro6ca66 , @logroll , @ThylacineAlive , @Hix , @snowleopard , @Newzooboy , @Patrick87 , @LaughingDove , @Sarus Crane , @Nick@Amsterdam , @taun , @KevinVar , @Giant Eland , @vogelcommando , @carlos55 , @TeaLovingDave , @devilfish , @Gust1 , @Kakapo , @Ituri , @geomorph , @Andrew_NZP , @GerbenElzinga , @alexkant , @keith8404 , @Dibatag , @Goura , @MagpieGoose and @Javan Rhino for their picture(s)

With 7 of the 12 African antelope tribes gone, over half of the African antelopes has already been covered. There are however several juicy species (complexes) left. The coming species which G&G have newly split are left:

NEOTRAGINI
Suni (Nesotragus moschatus)

AEPYCEROTINI
Common impala (Aepyceros melampus)

ALCELAPHINI
Bontebok/Blesbok (Damaliscus pygargus)
Topi/Tsessebe/Tiang/Korrigum (Damaliscus lunatus)
Hartebeest (Alcelaphus busephalus)
Common wildebeest (Connochaetes taurinus)

HIPPOTRAGINI
Sable antelope (Hippotragus niger)
Beisa oryx (Oryx beisa)

CEPHALOPHINI
Blue duiker (Philantomba monticola)
Common duiker (Sylvicapra grimmia)
White-bellied duiker (Cephalophus leucogaster)
Black-fronted duiker (Cephalophus nigrifrons)
Ogilby's duiker (Cephalophus ogilbyi)
Weyn's duiker (Cephalophus weynsi)
Yellow-backed duiker (Cephalophus silvicultor)
Bay duiker (Cephalophus dorsalis)

 

This is a fascinating resource thread, thank you for all the effort!

 

Yes. This is pretty much the free online version of "Bovids of the World" by Jose R. Castello. A great guide that's available on the iBooks store if anyone is interested. Great work on this list!

 

Happy to see a few of my photos show up on here

For the record the Greater Kudu at the San Diego Zoo Safari Park (and some others in the US) are of the nominate subspecies, and the Springbok at Fort Worth (as well as some other US zoos) are known nominate subspecies. Also the female klipspringer at Dallas and at least some individuals between the two San Diegos are stevensoni.

~Thylo

 

I assume they are what used to be the nominate, but what is now zambesiensis, or are they really from the Cape region?

 

Well obviously the thread can't compete with the book in the sense I am providing no info apart from diagnostics and taxonomy and Castello has a bit more pictures too though (though the pictures in this thread don't really suffer from all the issues with color and sun light...)

I pretty much disagree with most of the taxonomy used in the book though (I guess I made that point quite clear already). The funny thing is that the Castello book suffers from pretty bad editing, which is especially shown by the fact that in the text they often did not change "subspecies" with "species", even though the subspecies in question was split by Castello. So several G&G species are shown as species in the title, but referred to as subspecies in the main text.

Originally his book won't have looked much different from this thread when looking at the taxonomy used . First all the G&G splits were not included in the book, but given the huge number of errors, it seems a last minute change to incorporate all these splits without question, even though many are based on next to nothing....

 

Not entirely sure actually, but I will try to look into this further.

~Thylo