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

Glad to see this thread is back! Can't wait for the rest!

 

Common fallow deer

The Common or European fallow deer (Dama dama) is a medium-sized deer species from Anatolia and Southern Europe that has been spread by humans throughout much of Europe and countries worldwide. It is currently considered monotypic as the Persian fallow deer (Dama mesopotamica) has been split based on extensive morphological and genetic differences. Time of divergence between these two species has been estimated at 400.000 years ago.

Male

@Patrick87 , Zoo Eberswalde, Germany

Female

@Patrick87 , Zoo Eberswalde, Germany

Winter coat:

@Daniel Sörensen , Tiergarten Hannover, Germany


Persian fallow deer

The Mesopotamian or Persian fallow deer (Dama mesopotamica) is a critically endangered monotypic deer species which is currently restricted to two relict sites in Iran and has been re-introduced to other locations in Iran and Israel.

Male

@fofo , Tehran Zoo, Iran

Female

@nikola , Opel-Zoo Kronberg, Germany

 

Brow-antlered deer

The Eld's or Brow-antlered deer (Panolia eldii) is a large deer species from SE Asia. It has been placed in the genus Rucervus, but genetic research has shown it is actually a sister species to Pere-david's deer (Elaphurus davidianus) and they are the sister species of the genus Cervus. Alternatively they could be included in an extended Cervus genus but given the clear morphological and genetic differences that doesn't seem the best option. Traditionally three subspecies have been recognized and a fourth hainanus from Hainan Island has found little support in the literature:

P.e. eldii Sangai, NE India
P.e. thamin Thamin, Myanmar
P.e. siamensis Laos, Cambodia, Hainan Island, China and is possibly extinct in Vietnam & Thailand

G&G elevate all subspecies to species status

Sample sizes

No sample sizes for skins are given

Skulls
eldii 3
thamin 15-17
siamensis 3

Antler characteristics are ignored by G&G, except that eldii supposedly has short antlers

Skins

siamensis is described as being lighter in color and with spots visible in the median dorsal region (but see picture of female thamin below), thamin is described as being dark-brown in color. No description is given whatsoever for eldii except that it has bare pasterns.

Skulls

thamin and eldii are supposedly larger than siamensis and thamin is supposed to be relatively narrow-skulled. The data presented do however not support any of these conclusions, except when looking only at averages and ignoring the wide overlapping variation in all three taxa.

Additional data

In an earlier study Groves just notes that the different Eld's deer taxa are "extremely different", but he doesn't present the data on which this is based. There has been one study looking at genetic variation within Eld's deer (Balakrishnan et al. 2003) and they found all three subspecies to be genetically somewhat different, though the single eldii sample was nested within thamin. eldii is however ecologically somewhat different compared to siamensis and thamin as it prefers swampy areas and has thus adapted feet compared to the other two taxa. It has been noted that similarly as in Barasingha the adaptation to wetter habitats may be relatively plastic over evolutionary time and would not be a good indicator for speciation, though strict PSC taxonomists might disagree.

I am surprised there is no detailed description on the differences in horn shape between the taxa, as here there seem to be some clearer differences between the taxa.

Summarizing

Overall G&G do not present any data which would make the three different subspecies appropriate for consideration of species status. Additional future research would be necessary to see what the exact relationship is between eldii and thamin and whether eldii is just an ecomorph of thamin or whether it represents a distinct lineage. Extensive morphological research would also be welcome, though for now there is nothing clearly pointing in the way that there might be multiple species of Brow-antlered deer.

Panolia eldii eldii

@Chlidonias , Alipore Zoological Gardens, India

Panolia eldii thamin

@Maguari , Lisbon Zoo, Portugal

Female

@KevinB , Burgers' Zoo, Arnhem, Netherlands

Panolia eldii siamensis

@baboon , Guangzhou Zoological Gardens, China

Female:

@Himimomi , Guangzhou Zoological Gardens

References

Balakrishnan et al. (2006): https://repository.si.edu/bitstream/handle/10088/6138/Balakrishnan2003.pdf

Groves (2006): The genus Cervus in eastern Eurasia

 

During my research of this topic I have completely overlooked a recent paper studying mitochondrial dna of Red muntjac (Muntiacus muntiak s.l.) throughout its range:
Phylogeography of red muntjacs reveals three distinct mitochondrial lineages

They have found three clearly separate lineages. The oldest lineage is what has been normally described as malabaricus from Sri Lanka and the Western Ghats in SW India. They estimate that this population has split off from the other Red muntjac at around 1.5 mya. They also find a clear divide between northern (vaginalis) and southern (muntjak) Red muntjac and date the split at about 1.12 mya. These are split-off dates that are much older than in many currently recognized species such as Mesopotamian and Common fallow deer (dated at 400.000 years ago). They mention that there is likely a small hybridization zone between vaginalis and muntjak at the Isthmus of Kra, but give no evidence for this. Despite the clear differences found in their dataset, the authors do not argue that Red muntjac should be split in 3 species, but they ask for more research involving nuclear dna as well among other types of research. Regardless of the conservative approach that the authors take this is increased evidence for the recognition of multiple Red muntjac species after morphology and karyotype also provided some evidence. It also shows that even while the splits that G&G present are often somewhat outrageous given the evidence they provide, there is truth in some of their splits, and it seems to have got people working on some of their ideas. This is exactly what they envisaged and is much more helpful than carelessly excepting every split they propose (I am looking at you HMW...).

I also noted that I did not cover the remaining Muntjac species and I will get on with that soon, as with the genus Cervus, which is an absolute taxonomic nightmare/land-off-opportunity.

 

Port Lympne have a herd of about a dozen. I believe they were originally sourced from one of the Berlin zoos, probably East. I've seen them listed as branderi too, but think this may simply be incorrect. The wild population of branderi reached critically low levels in the 1970-80's with only around 70 left. I think its more likely that all/most swamp deer that have entered zoos are from the more numerous duvaucelii populations of Northern India.

 

I don't want to make too far a digression, but wearnt Aspinall's ssp. branderi imported in the same shipment as their Four honored horned antelope (Tetracerus quadricornis) from India? I think @TeaLovingDave told me this.

 

I think, along with Sus (scrofa, mostly) this is the genus I've been most looking forward to this entire thread. I've been looking into the red deer/wapiti complex lately and have pretty much just thrown my hands up in resignation. Best of luck, especially with all the wallichi/hanglu stuff.

 

I'd be interested to know exactly where Aspinall's Barasingha did come from, I can't remember why I think it was E. Berlin,(I thought I had read it in one of their reports but it was many years ago now) but any evidence they came from elsewhere would be interesting. Especially as they would represent a more recent import from the wild if they came from India.

 

I believe Berlin does indeed have some animals that were brought in from Berlin, but they also list themselves as keeping both the nominate and branderi (only one of the latter at this point).

~Thylo

 

I guess you mean Port Lympne not Berlin(third word..) I think I queried them once about this but got no response. It would be interesting to establish the facts here.

 

Oops! I meant to say I believe Port Lympne did get animals from Berlin, and PL lists themselves as having both. Thanks for the correction. But yes, some established facts would be nice.

~Thylo

 

Sika deer

The Sika deer (Cervus nippon) is a medium sized deer species from Eastern Asia and Japan. Taxonomy of this species has always been confusing given the large range and variation within its range. Over twenty different subspecies have been recognized and validity of several subspecies has
been called doubtful. The following subspecies have recently been accepted by various authors:

C.n. nippon South Japan
C.n. aplodontus N Honshu, Japan
C.n. centralis N & C Honshu, Japan
C.n. yesoensis Hokkaido, Japan
C.n. keramae Ryukyu Islands, Japan
C.n. mantchuricus Russian Far East, NE China
C.n. mandarinus N China (possibly extinct)
C.n. hortulorum N China (including dybowski)
C.n. grassianus Shansi, China (possibly extinct)
C.n. kopschi SE China
C.n. sichuanicus Sichuan & Gansu, China
C.n. taiouanus Taiwan
C.n. pseudaxis N Vietnam (possibly extinct in the wild)

G&G recognize 7 Sika deer species:

C. nippon (with keramae, yakushimae, mageshimae as subspecies and centralis as a synonym)
C. aplodontus (with yesoensis as a subspecies)
C. pulchellus (Tushima Island, normally considered as nippon)
C. hortulorum (including mantchuricus, grassianus as synonyms)
C. sichuanicus
C. taiouanus
C. pseudaxis (with mandarinus, kopschi as synonyms)

So confusing it is.

Sample sizes

Skins

No sample sizes for skins are given

Skulls (males/female)
yesoensis 4/0
aplodontus 4-6/0
nippon 1-5/0
nippon Goto Islands 9-10/8
nippon Kerama 0/0-2
yakushimae 1-2/0
mageshimae 0-1/0
pulchellus 1-4/2
Beijing 0-1/0
hortulorum 0-1/3
sichuanicus 0-1/0-2
kopschi 14-16/6
grassianus 0-1/0
pseudaxis 2-3/0
Solo Island, Phillipines 1 (introduced population)

Sample sizes are thus mostly extremely limited and almost completely lacking for females. No
samples for taiouanus are given at all.

Skins

aplodontus is described as having a very large neck mane, yellowish white hair around metatarsal
glands, lighter-colored face, darker winter coat and a constant presence of a row of spots along
median dorsal line. Contrary nippon should have a small neck mane, only indistinct spotting along
dorsal line, a large rump-patch, dark metatarsal glands surrounded by light grey fur. The only skin
difference listed for pulchellus is a whitish belly. hortulorum is said to have a reduced or absent neck
mane, light red-brown to grey winter coat and large rump-patch. No mane is supposedly present in
sichuanicus, but dorsal stripe is distinct with row of white spots on either side. taiouanus is
described as clearly spotted in winter, in summer it is bright chestnut with prominent dorsal stripe
and males are strongly maned. pseudaxis is described as somewhat resembling taiouanus with
bright summer coat, gray to black winter coat, spotted in both seasons, underside light brown. G&G also give differences in tail length, but do only state the mean of measurements, so it is impossible to see whether this characteristic is meaningful when taking into account variation.

Skulls & antlers

nippon, from S Japan is described as being bigger than aplodontus from N Japan and this is generally
true and despite a limited sample size, there is hardly any or no overlap in several skull
measurements. pulchellus overlaps widely with nippon in skull measurements, but is said to have a
long basal segment to the antlers. sichuanicus is described as being large, but there is wide overlap
in most measures with pseudaxis and hortulorum. When comparing both Japanese taxa nippon and
aplodontus with mainland species there is wide overlap in several measurements. taiouanus antlers
are being described as rather weak but the only one to have pinkish velvet, as opposed to black in
the other taxa.

Additional data

Sika deer have been the focus of extensive genetic research over the past 20 years, but I will focus
here on the most recent research, including the largest geographic sample and the largest number of
base pairs (all mtdna). Ba et al. (2012) partly took the original data from earlier studies and they
found four distinct clades of Sika deer, some further subdivided. They estimate that at about 1.72-
2.03 mya Sika deer from northern Japan (aplodontus) diverged from the other taxa. At about 1.45-
1.72 mya Sika deer from southern Japan diverged from the mainland. Afterwards Sika deer from
Yakushima and Mageshima diverged from the other S Japanese Sika deer (nippon). The split
between the northern mainland subspecies (hortulorum, mantchuricus) and the Southern mainland
+ Taiwan subspecies (pseudaxis, taiouanus, sichuanicus, kopschi) is dated at 0.18-1.08 mya. Despite
these long periods of separation genetic divergence is estimated between 0.5 and 4.6 percent
between these four major groups. Only the Northern Japanese sika deer (aplodontus) have a genetic
divergence of over 4% with the mainland taxa. Mainland taxa and Southern Japanese Sika deer
(nippon) have a genetic divergence of about 3.7-3.9%. The two mainland groupings have a genetic divergence of roughly 2.5%. These genetically identified groups are also supported by Yang et al.
(2012) and more or less agree with earlier genetic studies.

To make matters more complicated Sika deer have also been introduced in several parts of the worlds. Sika deer in Central Europe belong to both the Northern and Southern Japanese stock, as well as from the northern mainland population (Pitra et al. 2005; Barancekova et al. 2012).

According to Takatsuki (2009) the Northern Japanese and Southern Japanese sika deer also differ in their feeding habits, with northern being mainly grazers and southern being mainly browsers.

Summarizing

Overall based on the limited data presented by G&G there is little reason to follow their proposed
species delimitations. Combined with genetic work there is however strong support for the recognition of 4 major groups of Sika deer, two in Japan and two on the mainland + Taiwan. These 7 species proposed by G&G fit in these four groups, but are seriously over-split based on hardly any data. Despite the longtime of divergence the genetic differentiation between these four major groups is relatively small, but may in some case approach species status. Karyotype seems to be mostly 2n=66 though and there is no research yet presenting results with nuclear dna as far as I am aware. This should however be a priority to assess whether the four main groups of Sika deer are better treated as distinct species than as the subspecies they are now.

To summarize the provisional Sika deer grouping would look like this:

C. (n.) nippon S Japan (including pulchellus, yakushimae, mageshimae, keramae)
C. (n.) aplodontus N Japan (including yesoensis, centralis)
C. (n.) hortulorum Far East of Russia, N China (including manchuricus)
C. (n.) pseudaxis S China, Taiwan, N Vietnam (including taiouanus, sichuanicus, kopschi)

The status of subspecies within these groups is still unclear though.

Note that the taxonomy of Sika deer as presented in the zootierliste is a mess, it is based on a combination of sources without cross-checking

C. (n.) nippon

@Swedish Zoo Fan , Ueno Zoo,, Tokyo, Japan

C. (n.) nippon yakushimae

@devilfish , Inokashira Park Zoo, Japan

C. (n.) hortulorum

@Al , Zoo Antwerpen, Belgium


@Arek , Zoo Wroclaw, Poland

C. (n.) hortulorum mantchuricus (winter coat)

@mhale , ZSL Whipsnade Zoo, UK

Summer coat:

@Maguari , Woburn Safari Park, UK

C. (n.) pseudaxis pseudaxis

@Patrick87 , Tierpark Berlin, Germany


@vogelcommando , Safaripark Beekse Bergen, Hilvarenbeek, Netherlands

C. (n.) pseudaxis taiouanus

@aardvark250 , Taipei Zoo, Taiwan


@Giant Eland , Taipei Zoo, Taiwan

possibly C. (n.) pseudaxis kopschi

@YuanChang , Fuzhou Zoo, China

No pictures of Northern Japanese Sika (aplodontus) have been uploaded to the gallery. The status of "Japanese sika" in European and US zoos is unclear.

References

Ba et al. (2012)
https://www.researchgate.net/profile/Hx_Ba/publication/257070414_Classification_and_phylogeny
_of_sika_deer_Cervus_nippon_subspecies_based_on_the_mitochondrial_control_region_DNA_seq
uence_using_an_extended_sample_set/links/54f66b0b0cf21d8b8a5c6aaa/Classification-and-
phylogeny-of-sika-deer-Cervus-nippon-subspecies-based-on-the-mitochondrial-control-region-DNA-
sequence-using-an-extended-sample-set.pdf

Barancekova et al. (2012): The Ecological Society of Japan Journals

Pitra et al. (2005): Tracing the genetic roots of the sika deer Cervus nippon naturalized in Germany and Austria

Takatsuki (2009): Geographical Variations in Food Habits of Sika Deer: The Northern Grazer vs. the Southern Browser

Yang et al. (2012) https://www.tandfonline.com/doi/abs/10.1080/00222933.2012.693959

 

Red deer

The Red deer (Cervus elaphus) is a large species of deer from Eurasia and a small part of N Africa. Red deer have been considered conspecific with Wapiti (Cervus canadensis), from E Asia and N America, but Wapiti is actually the sister species to Sika deer instead of Red deer and will be discussed separately. Red deer have a wide distribution and a complicated evolutionary history, so a large number of different subspecies have been described and have been recognized recently:

C.e. elaphus Ireland, UK, W and N Europe (including scotticus, hippelaphus and atlanticus)
C.e. hispanicus Iberian Peninsula
C.e. italiensis Currently restricted to 1 location in the Po delta
C.e. corsicanus Sardinia & Corsica
C.e. barbarus E Algeria, W Tunisia
C.e. pannoniensis parts of C and SE Europe (also treated as montanus, includes some populations regarded as hippelaphus)
C.e. maral Turkey & Caucasus
C.e. brauneri Crimea
C.e. yarkandensis Tarim Basin, China
C.e. bactrianus N Afghanistan, S Uzbekistan, S Turkmenistan
C.e. hanglu Kashmir Vale, India

G&G recognize 7 species of Red deer:
C. elaphus (including hispanicus, italicus)
C. pannoniensis
C. corsicanus (including barbarus)
C. maral
C. yarkandensis
C. bactrianus
C. hanglu

Sample size

No sample sizes are given for skins, skulls or antlers. Additionally no skull or antler data are presented so it is impossible to verify any of their claims.

Skins

No data is given on how most species differ in colouration. yarkandensis and bactrianus are regarded as very similar, but it is not said how they differ from the others. hanglu is morphologically apparently more similar to deer from E Asia (canadensis group). maral is also said to be quite distinct in color, spotting and mane, but the reference to this is not available online.

Skulls & antlers

No data on skulls are presented or mentioned. Nor is a comparison of the different antlers given between the taxa.

Additional data

G&G rely very strongly on a growing body of genetic work that has looked into genetics of this complex group (and presumably on some unpresented analyses).

Genetically there is a clear split between yarkandensis, hanglu and bactrianus and the other Red deer. Lorenzini & Garofalo (2015) estimate a split about 4 million years ago. hanglu from Kashmir is distinct, but it is unclear whether bactrianus and yarkandensis are valid taxa in their analyses or whether they should be lumped. These taxa are very similar and separated by the Tian Shan mountain range. Ludt et al. (2004) treat them as subspecies, but as only 1 sample for each was present their monophyly cannot be confirmed. Kuznetsova et al. (2012) can also not resolve this. There is however broad agreement to treat these three taxa as Cervus hanglu.

The second Red deer group is however more difficult with a larger number of taxa described and many (historic) translocations. There is however broad agreement that there are several distinct taxa, mainly based on different glacial refugia:

- elaphus which spread from a refugium in SW Europe/Iberian Peninsula throughout W and many parts of N Europe

- pannoniensis which spread from a refugium in the Balkans throughout C and SE Europe

- italiensis a recently described taxa from a refugium in the Po Delta Italy (Zachos et al. 2015) and is closer related to pannoniensis and maral then to elaphus

- corsicanus Including deer from N Africa & Corsica + Sardinia

- maral from Turkey & the Caucasus.

barbarus is thus lumped with corsicanus. The deer from Corsica and Sardinia are however probably Early Holocene human introductions and have since become smaller. Based on genetic data the source is thus likely N Africa.

hispanicus from the Iberian Peninsula has often been lumped with elaphus. Rey-Iglesia et al. (2017) and Fernandez-Garcia et al. (2014) do however find strong indication that there is a separate lineage of Red deer on the Iberian peninsula (though there have been translocations from other parts of Europe, so this original population is threatened). This would mean hispanicus would be a valid subspecies. Queiros et al. (2015) hypothesize that the distinction of hispanicus means that it is likely elaphus has a refugium N of the Pyrenees, either SW France, the continental shelf or S Ireland. There is no genetical support for the subspecies scotticus, hippelaphus and atlanticus, which makes sense as these areas have only been recently colonized after the last glaciation event.

The challenge is now to assess what this geographical structuring means, are these taxa better defined as subspecies or as species. There are indications of a mixing zone between pannoniensis and elaphus in parts of Central Europe. The date of divergence between the main Red deer groups in Europe is given at about 200.000 - 300.000 years by Skog et al. (2009) and genetic divergence between these European groups is rather small. Time of divergence between The European red deer and barbarus is estimated at about 1.1 mya by Pitra et al. (2004). maral is estimated to have diverged 1.4-1.7 mya ago from barbarus+elaphus+pannoniensis. Lorenzini & Garofalo (2015) do however put the split between barbarus and the European red deer at 2.4 mya (with a large error margin from 0.9-4.5 mya). Meiri et al. (2017) do however find that maral, pannoniensis and italicus are closely related with elaphus and hispanicus are sister to these taxa. So there is still some unclarity about divergence times and exact relationships. Unfortunately there seems to be no data on potential differences in karyotype of differences in nuclear dna.

Summarizing

There is clear evidence that at least two species of Red deer should be recognized: C. elaphus from Europe, N Africa and Anatolia and C. hanglu from Central Asia. It remains somewhat unclear whether maral or corsicanus (corsicanus precedes barbarus so the Red deer of N Africa should be collared barbarus) would also deserve species status and how they are exactly related to other Red deer. Provisionally it would make sense to recognize two species, each with a number of subspecies. This will however not be the final classification but is a compromise based on current evidence:

C. hanglu
C.h. hanglu
C.h. yarkandensis
C.h. bactrianus

C. elaphus
C.e. elaphus
C.e. ibericus
C.e. pannoniensis
C.e. italicus
C.e. maral
C.e. corsicanus (includes barbarus)

C.h. bactrianus

@gulogulogulo , Highland Wildlife Park, Kingussie, UK


@Kudu21 , The Wilds, USA

C.e. elaphus

@Kibathewolf , Orostrand, Denmark


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

C.e. hispanicus (note that elaphus also occur wild in Spain, so these pictures may show hybrids)

@Giant Eland , Safari Madrid, Spain


@Maguari , Zoo-Aquarium Madrid, Spain

C.e. corsicanus

@thor , San Diego Zoo Safari Park, USA


@ThylacineAlive , Tierpark Berlin, Germany

C.e. pannoniensis

@alexkant , Ploiesti Zoo, Romania

C.e. maral

@fofo , Iran in the wild

No pictures of hanglu, yarkandensis or italicus have been uploaded to the gallery.

References

Fernandez-Garcia et al. (2014): Mitochondrial D-loop phylogeny signals two native Iberian red deer ( Cervus elaphus) Lineages genetically different to Western and Eastern European red deer and infers human-mediated translocations

Lorenzini & Garofalo (2015): Error - Cookies Turned Off

Ludt et al. (2004): https://www.researchgate.net/profil...phylogeography-of-red-deer-Cervus-elaphus.pdf

Meiri et al. (2017): Error - Cookies Turned Off

Pitra et al. (2004): https://s3.amazonaws.com/academia.edu.documents/32525637/Pitra_et_al_2004_Mol_Phylogenet_Evol_Evolution_and_phylogeny_of_old_world_deer.pdf?response-content-disposition=inline; filename=Evolution_and_phylogeny_of_old_world_dee.pdf&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Credential=AKIAIWOWYYGZ2Y53UL3A/20190628/us-east-1/s3/aws4_request&X-Amz-Date=20190628T165408Z&X-Amz-Expires=3600&X-Amz-SignedHeaders=host&X-Amz-Signature=5989c1316cc3faddc35205692e8b1400bb1db8f13456a9e75403fb3914676c9b

Queiros et al. (2015): Red deer in Iberia: Molecular ecological studies in a southern refugium and inferences on European postglacial colonization history
Rey-Iglesia et al. (2017): Error - Cookies Turned Off

Skog et al. (2009): Error - Cookies Turned Off

Zachos et al. (2015): https://www.tandfonline.com/doi/full/10.1080/11250003.2014.895060

 

@lintworm Kudos (or should I say Kudus) on this fantastic work!

Admittedly I'm not quite clear exactly how MacNeill's or Sichuan Deer should be categorized, but from the scientific names offered on zootierliste:

Cervus wallichii hanglu
(Syn.: Cervus elaphus hanglu)
(Syn.: Cervus hanglu)

perhaps my photos here: 2013: Shadow Nursery - ZooChat
& 2013: Shadow Nursery - ZooChat
would qualify for Cervus hanglu hanglu- not yet represented here.

 

Zootierliste is outdated, as it is clear now that hanglu belongs to the elaphus group, despite morphological characters that look more like the canadensis group. Red deer from E China belong to the canadensis group, whether multiple species should/could be recognized is something for the next post.

 

Thanks for clearing that up. I've always been confused about the taxonomy of that species/subspecies. Looking forward to the next post!

 

I've since uploaded photos of smithemani taken in Spain:
Black Lechwe (Kobus leche smithemani) - ZooChat

@lintworm also Gazella dorcas isabella
Isabelline gazelle (Gazella isabella) - ZooChat

 

There has been new research published on Bushbuck taxonomy and it all but confirms the split between scriptus and sylvaticus:

https://www.sciencedirect.com/science/article/pii/S1055790318304354

This study found differences in the number of chromosomes between both taxa as well as clear differences in nuclear as well as mitochondrial dna. Mitochondrial dna places scriptus as sister to Nyala, but nuclear dna doesn't, which is likely because of ancient hybridization events between a an extinct close relative of Nyala and ancient scriptus populations.

The study also finds Ethiopian highland bushbuck meneliki to be well nested in sylvaticus. There seems thus to be a very strong case now to recognize 2 species of bushbuck, each with a number of subspecies (could maybe be called species under extremely strict PSC criteria).