Capricious subspeciation in the plains zebra, part 2: introducing the endangered Gorongosa zebra, Equus quagga foai

(writing in progress) 

...continued from https://www.inaturalist.org/journal/milewski/56214-capricious-subspeciation-in-the-plains-zebra-part-1#

The plains zebra (Equus quagga) was first discovered by Science in the eighteenth century. From the start, the conservation of the many and various subspecies of this widespread species has been hindered by a labelling failure.

This problem has continued until today - despite the sophisticated methods of modern taxonomy.

Equus quagga quagga was originally abundant and fairly widespread in South Africa. However, it was unwittingly hunted to extinction, because of an assurance that there remained plenty of 'kwaggas' farther north, across the Vaal River (https://en.wikipedia.org/wiki/Vaal_River). Unbeknownst to landowners, the 'kwagga' north of what is now Free State province (https://en.wikipedia.org/wiki/Free_State_(province)) was a different subspecies - so different, indeed, that it might as well be a different species.

Had the extinct quagga (https://en.wikipedia.org/wiki/Quagga) been given a more precise name, this nomenclatural confusion - and the biological tragedy caused by it - might have been avoided.

This is, indeed, why the quagga (E. q. quagga) was allowed to slip into extinction when other sympatric ungulates (e.g. black wildebeest) were saved. Because ‘kwaggas’ were still known to be abundant farther north in South Africa, nobody saw E. q. quagga for the distinctive form that it was, until it was too late. Actually, much too late (it was decades after extinction that anyone woke up to the loss).

One particular form has posed a taxonomic puzzle for many years.

I have, for the last half-century, recognised the intensely-striped zebra of Gorongosa National Park (https://hfmoconservationandscience.weebly.com/education-ecology-and-conservation/the-history-and-biodiversity-of-the-gorongosa-national-park and https://en.wikipedia.org/wiki/Gorongosa_National_Park) as something distinct.

Although Equus quagga crawshayi, of eastern Zambia, Malawi, and northern Mozambique has narrow stripes, the Gorongosa zebra has stripes that are even narrower, while retaining bold definition (https://www.flickr.com/photos/zimbart/38718489444).

Although fairly frequently photographed in Gorongosa National Park in the 'sixties and 'seventies, the Gorongosa zebra was virtually exterminated in that Park during the Mozambican Civil War. (https://en.wikipedia.org/wiki/Mozambican_Civil_War).

However, until now I have assumed that the Gorongosa zebra has never been given a scientific name that we can use to designate it as a particular subspecies of the plains zebra (Equus quagga).
 
As you can understand, it’s hard to push for the conservation of a critically endangered taxon that has no name. People tend not to take it seriously if it’s not labelled clearly.
 
Prazak and Trouessart in 1899 (Bull.Mus.Paris, 5:350, 1899) described an extremely intensely striped form from Mozambique as ‘foai’, so I think there’s a good chance that the Mozambique zebra can validly be called Equus quagga foai.

The type locality of what they described as a distinct species, Equus foai, is "the mountainous district of Angoniland, on the north bank of the Zambezi opposite Teti". See the location in https://en.wikipedia.org/wiki/Tete,_Mozambique.

Lydekker (1916, Catalogue of Ungulate Mammals in the British Museum) says that it is distinguished by "the larger number of main stripes on the body and hindquarters, and the absence of any backward bending (except in the last of the series) of the body stripes as they approach the dorsal stripe, to which they run approximately at right angles. In this respect Foa's zebra approximates to the true zebra and Grévy's zebra, from both of which it differs by the stripes on the hindquarters adjacent to the dorsal stripe running parallel with the latter in the direction of the tail…" 
 
The hide specimen shown below certainly belongs to the Mozambique zebra, and it’s the first such picture I’ve ever seen. It would make a good illustration for any item we post on EB on the topic of the Mozambique zebra, not so?

Something I have not noticed before: the striping is so intense on the posterior part of the dorsal area, or the anterior rump, that it forms a virtual spotting of pale on a ‘ground-colour’ of dark! I’ve never seen that in any other type of zebra. It is a noteworthy feature because it

  • transcends striping to the point of qualifying as spotting, and
  • inverts the normal relationship of dark and pale in this species.

http://www.planet-mammiferes.org/Photos/Ongule/Equides/EquuBFoa.jpg

http://www.planet-mammiferes.org/drupal/en/node/64?choixzone=&input2=Eastern+Africa&plus=0&fin=3
  
The hide specimen shown below certainly belongs to the Gorongosa zebra, and it is the first such picture I have ever seen.
 
(writing in progress)

Posted on July 13, 2022 10:29 AM by milewski milewski

Comments

For years, I have placed ‘crawshayi’ in apostrophes. This is because a) I have seen the form in e.g. in Luangwa to be intermediate between the Mozambique zebra and Grant’s zebra (E. q. boehmi) of western Zambia, and b) there is much intergradation among subspecies in the whole Zimbabwe/Zambia/Mozambique area (e.g. Paul Dutton has persistently referred to the Gorongosa zebra as Selous’ zebra, Equus quagga selousi).

Posted by milewski almost 2 years ago

Photo showing back-of-ear flag (posterior auricular flag) of plains zebra in Equus quagga boehmi in Ngorongoro caldera:
http://l7.alamy.com/zooms/cec628dfcf574a7093ea56ead8b2e4c8/three-zebras-turn-their-backs-to-run-away-in-the-ngorongoro-crater-b8kkad.jpg.

I suspect that the dark markings on the back-of-ear are more extensive than this in Equus quagga foai.

Posted by milewski almost 2 years ago

This is a great emphasis on phenotypical distinctions within the Plains Zebra, it’s remarkable the Gorongosa population has survived for us to see the features present. However, the convention of Equus quagga being of many subspecies is far from settled, as there exists studies that conflict with the respective convention. There is also ample evidence of a bottleneck ("A rapid loss of stripes: The evolutionary history of the extinct quagga") in a prehistoric population of the southern plains zebra grouping that led to the extinct quagga. From a viewpoint concerning spread of traits derived from a minimized population, it (the bottleneck & it’s effects) possibly led to the unique ecophenotype present in the extinct nominate race of Equus quagga.
link to the study relating to the Quagga : https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1617154/

Posted by paradoxornithidae over 1 year ago

This Nature Magazine article details the divergence time for the extant populations of Equus quagga, & apparently, according to some taxonomic authorities, it dissolves the subspecies differentiations currently recognized.
https://www.nature.com/articles/s41559-017-0453-7

Posted by paradoxornithidae over 1 year ago

This also has implications in the conservation of these taxa, would it be better to admix other plains zebra with the Gorongosa zebra?

Posted by paradoxornithidae over 1 year ago

@milewski What is your view on the validity of the currently delineated subspecies of Equus quagga?

Posted by paradoxornithidae over 1 year ago

@paradoxornithidae

I discussed this topic in detail with Colin Groves, before he died several years ago.

This is my current opinion.

Firstly, it is difficult to treat the plains zebra taxonomically, because it is, in several ways, odd in its differentiation/speciation/subspeciation.

However, assuming that this is one widespread species (Equus quagga), I would list its subspecies as follows (excluding extinct possible subspecies in Africa northwest of the current range).

quagga extinct

burchellii eastern Free State to Zululand

unnamed ssp., phenotypically similar to burchellii but disjunct northern Namibia

chapmani Mpumalanga to Okavango and Zambezi mouth

foai Gorongosa, southern Malawi, and possibly up to Northern Mozambique

crawshayi Luangwa Valley, Nyika Plateau, and southern Tanzania

boehmi western Zambia all the way to central Kenya

'maneless' ssp. in Uganda and southern Sudan

'maneless' ssp. in Somalia, disjunct from the last-mentioned (intergraded with boehmi in Tsavo)

subspecies in Ethiopia (not maneless, and featuring more shadow-striping than boehmi)

This totals up to ten sspp.

Posted by milewski over 1 year ago

@milewski thanks for the detailed response

Posted by paradoxornithidae over 1 year ago
Posted by milewski over 1 year ago

@paradoxornithidae

In https://www.youtube.com/watch?v=f_r9oQFjJUs, you can get a glimpse of the difference between the form of Equus quagga in Tsavo East National Park in Kenya, and the more familiar form in e.g. the Serengeti. The individual at the rear of this small group is, I think, adult male, and you should be able to see its 'manelessness'. I suspect that this population, in Tsavo East, is intergraded between ssp. boehmi and the Somali ssp. (which may already be extinct in pure form).

Posted by milewski over 1 year ago

interesting, thanks

Posted by paradoxornithidae over 1 year ago

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