DANIELL'S QUAGGA AND WARD'S ZEBRA - ANOTHER TWO STRIPED CURIOSITIES OF THE EQUINE KIND

 

Daniell's quagga (left) and Ward's zebra (right) (public domain)

Following on from my previous ShukerNature article concerning the beautiful but long-forgotten isabella quagga (click here to access it), here are another two eyecatching but exceedingly obscure striped curiosities of the equine kind, retrieved from the annals of zoological history.

 

DANIELL'S QUAGGA – THE MOST EXTREME QUAGGA OF ALL?

Yes indeed, this particular quagga specimen is so extreme that it makes even the isabella quagga seem positively commonplace by comparison!

The specimen in question is a truly remarkable beast known as Daniell's quagga, after the artist Samuel Daniell (1775-1811), who produced a very handsome aquatint of it in 1804 for his African Scenery and Animals at the Cape of Good Hope two-part series (1804-1805). He based it upon this quagga form's only known specimen, which had been shot in southern Africa's so-called Square Mountains (currently unidentified by me) during 1801, but whose skin was not retained.

 

Daniell's quagga, painted by Samuel Daniell as it would have looked when alive in 1801 (public domain)

What was so extraordinary about it, as readily seen in Daniell's painting, is that this quagga specimen had exceptionally reduced striping. Indeed, the latter markings were confined almost entirely to the sides of the animal's neck, with only a few very faint lines upon its throat and shoulders, and none at all upon its torso. (True, I have seen paintings of certain other quagga specimens with stripeless torsos, but their throat and shoulders in addition to their neck all bore distinct, conspicuous stripes.) It also had a noticeably large head.

As with the isabella quagga, this specimen was initially deemed to represent a new zebra species, dubbed Daniell's quagga, and was accordingly given the species name danielli. However, and once again like its isabelline relative, Daniell's quagga was later subsumed into the plains zebra species Equus quagga as merely a non-taxonomic freak individual.

 

WARD'S ZEBRA – A ZEBRA CROSSING IN EVERY SENSE!

Ward's zebra is a distinctively-striped, long-eared interspecific hybrid resulting from matings between plains zebras E. quagga and mountain zebras E. zebra that was first brought to scientific attention in 1904 via a Proceedings of the Zoological Society of London report by British zoologist Prof. J.C. Ewart. In his report, Ewart stated that some years previously he had been presented with a taxiderm zebra specimen, the subject of his report, by Rowland Ward, who was a very famous London-based taxidermist at that time. Ewart had subsequently donated it to Edinburgh's Royal Scottish Museum (now part of the National Museum of Scotland).

According to Ward, the specimen had originally been "traded out of Somaliland", Somaliland nowadays being recognized as a region within Somalia. However, Ewart speculated that its kind "probably inhabits part of the area between the upper reaches of the Tana River and Lake Rudolf [later renamed Lake Turkana]", in Kenya.

 

Ward's zebra - two views of Ewart's erstwhile taxiderm hybrid specimen, from his 1904 PZSL report (public domain)

Ewart was struck by the specimen's overall similarities to South Africa's Cape mountain zebra (E. z. zebra; Hartmann's mountain zebra E. z. hartmannae occurs in Namibia and Angola), but also noting in detail various differences in its striping, as well as its very long ears. Clearly not suspecting its hybrid nature, Ewart concluded his report by suggesting that it may constitute a new form of Kenyan plains zebra, duly dubbing it Ward's zebra in honour of its procurer, which "is adapted to a habitat similar to that of the mountain zebra", i.e. an example of convergent evolution.

In 1910, moreover, Ward's zebra was formally named Equus wardi, but its hybrid status was revealed via the discovery that specimens of this zebra form had been obtained repeatedly in the Jardin des Plantes, Paris, around 1900. And in 1915, a male specimen was obtained at London Zoo. Indeed, some authorities have opined that Ewart's specimen had itself probably been bred in a menagerie, rather than originating from either the wilds of Somaliland or of Kenya.

 

Vintage engraving of the Cape mountain zebra, 1830 (public domain)

 

THE ISABELLA QUAGGA - A LONG-LOST, LONG-FORGOTTEN EQUINE ENIGMA

 

A beautiful vintage (1800s) full-colour illustration of the unique isabella quagga (public domain)

The quagga Equus quagga quagga is nowadays famous for two very different reasons. Firstly: it is – or was – the only semi-striped form of zebra, its striping being confined to its foreparts. Secondly: although once common in its South African veldt habitat, it was hunted into extinction there during the second half of the 19^th Century, with the very last captive specimen's death in Amsterdam Zoo on 12 August 1883 marking the tragic disappearance of this highly distinctive equid from the face of our planet – though The Quagga Project continues its aim to recreate this vanished creature's characteristic phenotype (external appearance) via back-breeding, using striping-deplete specimens of other Equus quagga subspecies to produce quagga facsimiles.

Speaking of which: today, the quagga is classed as a subspecies of the plains zebra Equus quagga, but back in the mid-1800s when still very much alive it was deemed to be a valid, distinct species in its own right, and was dubbed Hippotigris quacka (hippotigris being the name given to zebras by ancient scholars who believed these exotic-looking striped equids to be the product of matings between horses and tigers!) – see later for further taxonomic details. But that is not all.

 

One of five precious photographs of an adult quagga mare living at London Zoo from 15 March 1851 until her death there on 15 July 1872 – these are the only known photos of a live quagga (click here for more details concerning this quagga quintet)

For a time during that same period, a second, very remarkable quagga species was also recognized, despite being known from just a single specimen – a poorly-preserved skin formerly held at the British Museum in London. This unique, extraordinary-looking animal became known as the isabella quagga, but today the skin is long lost and the isabella quagga itself is long forgotten. Consequently, I felt that what (very) little is known about this beautiful if baffling enigma of an equid richly deserved to be collated and presented in article form in order for modern-day readers to become aware of its erstwhile existence. So here is the hitherto-obscure history of the long-overlooked isabella quagga – a ShukerNature exclusive.

I first learned of the isabella quagga Hippotigris isabellinus many years ago, when I chanced upon the following previously-obscure yet fascinating excerpt from a quagga-themed communication by famous British zoologist Richard Lydekker that had been published by the scientific journal Nature on 10 January 1901. The excerpt alluded to a supposedly separate, second species of quagga, again extinct:

...the British Museum formerly had the skin of a young quagga, in very bad condition, which was presented by the traveller William Burchell [after whom Burchell's zebra is named], and was subsequently described by Hamilton Smith as a distinct species, under the name of Hippotigris isabellinus.

Two points to note here. Firstly: the above-mentioned Hamilton Smith was Charles Hamilton Smith (1776–1859), a lieutenant-colonel in the British Army. He was also a naturalist who scientifically described and named several equine species and subspecies. In two 1841-published tomes referred to later here, he dubbed this enigmatic animal the isabella quagga. Secondly: whereas all zebra species and subspecies are nowadays housed in the genus Equus (alongside horses and asses), back in Lt-Col. Hamilton Smith's time several were housed in their own separate genus, Hippotigris, including the normal quagga, which was formally deemed back them to be a valid species in its own right (rather than merely a subspecies of the plains sebra, as it is classified today) and was duly known as Hippotigris quacka. 

 

Late 1800s chromolithograph from my personal collection, depicting a normal quagga with a bushbuck and a gnu (public domain)

Lydekker's communication then continued with the following text, but it is unclear whether this text was still referring to the isabella quagga or (as I suspect) had returned to the communication's primary subject, the normal quagga:

Apparently London museums possess no other relics of this lost species, of which, however, we believe there is a specimen in the museum at Edinburgh. As the animal yielded no trophies worthy the attention of the sportsman, it is unlikely that there are any specimens in private collections, unless, perchance, a skull or two may be in existence.

The remaining text in Lydekker's communication unequivocally referred to the normal quagga, so it needn't be quoted here.

What exactly was the isabella quagga, I wondered, when I first began researching this curious creature, and what did it even look like, bearing in mind that Lydekker provided no description of it in his communication and the British Museum no longer has it?

Back in pre-internet times, it was by no means easy to research anything as unimaginably obscure as the isabella quagga, so after various attemptss to solicit more information concerning it all proved futile, I placed Lydekker's intriguing communication on file and directed my attention to other subjects. Notwithstanding these failures, however, I never forgot about it, so when I was checking some details recently while completing some other researches and noticed it again, still on file, I decided to reinvestigate its elusive subject, but now assisted enormously by the vast wealth of data readily accessible online. And this time, finally, I was successful, as now revealed.

Originally, my only clue had lain in its moniker. For in this instance, isabella refers not to a woman's name but instead to a colour, known in full as isabelline, and which constitutes this mystery quagga's species name, isabellinus. It is variously defined as pale grey-yellow, pale fawn, pale cream-brown or parchment colour, and is primarily utilised in relation to mammalian coat colour and bird plumage.

Presumably, therefore, I mused, this shade was the background colouration of the coat of this unique specimen (a male, incidentally), meaning, if so, that it was paler in appearance than normal quaggas and probably with fainter stripes too. Whether such a difference warranted Hamilton Smith naming it as a separate species, however, when it was surely nothing more than an aberrantly pallid (possibly leucistic?) specimen of the normal quagga (see later), was another matter.

 

The pale-coloured engraving of the isabella quagga from Hamilton Smith's two 1841 tomes (public domain)

During my recent researches, I uncovered two beautiful vintage illustrations depicting the isabella quagga, both of which represent it in the living state. One of these illustrations is a hand-coloured engraving in very pale shades with minimal background colouration. The other illustration is in full-colour, so it is much more vibrant.

I traced the pale engraving back to a couple of tomes from 1841, which upon close examination turned out to be identical in content but bearing different titles. One is entitled Horses, and constitutes Volume 20 of the massive 40-volume series edited by Sir William Jardine and entitled The Naturalist's Library. The other tome is exactly the same but is retitled as The Natural History of the Horse and constitutes a stand-alone volume. In both tomes, the author is given as Charles Hamilton Smith, and a concise section documenting what he specifically refers to as the isabella quagga is included, containing the pale engraving of this specimen. In both tomes, it is designated as Plate 25, and is credited to Hamilton Smith.

In his duplicated 1841 tomes, Hamilton Smith began his brief coverage of the isabella quagga (pp. 332-334, and which constitutes this claimed species' formal scientific description and naming) by stating that although this animal's body shape (including its head) compared closely with that of the normal quagga, he had separated it from the latter equid because it differed by virtue of its smaller size (barely 10 hands, i.e. 40 in, tall) and even more so by the forms and colour of its stripes.

He then referred to an unidentified equid seen by travelling French naturalist François Le Vaillant (1753-1824), presumably in South Africa's Cape as this is where he had spent time collecting animal specimens, and which he'd named the zebre but was apparently different from those zebras already known from there. Some zoological authorities, including Dutch zoologist Coenraad Temminck (whose father was Le Vaillant's employer) had considered the isabella quagga to be Le Vaillant's zebre, but Hamilton Smith disagreed with their opinion.

The remainder of Hamilton Smith's account consisted of a verbal description of the isabella quagga skin (augmenting the engraving of this animal portrayed in the living state), which included his belief that it was an adult rather than a juvenile specimen despite its small size, and was not albinistic. Conversely, when concluding his account by mentioning that a Dr Leach had believed the skin (which still existed at the British Museum at this time) to have originally come from the Cape, he conceded that Leach had considered its pale colouration, especially its white stripes, to be due to the animal's 'nonage' (young age).

Moreover, it should be noted here that back in Hamilton Smith's time, there was a somewhat naïve but very prevalent tendency among taxonomists to over-emphasise the significance of individual variation within species, leading to the splitting off and naming of many spurious animal species that in reality were nothing more than freakishly-coloured/patterned individuals of already known, confirmed species. Eventually, however, such shortcomings were rectified by lumping these unsubstantiated species back together – as happened with the isabella quagga, subsequently being subsumed by zoologists into the normal quagga species (now subspecies).

 

Hamilton Smith's undated full-colour wtarcolour painting of the isabella quagga (public domain)

As for the full-colour isabella quagga illustration: it is an undated watercolour painting, again by Hamilton Smith, and is contained with various others of his watercolours in an unpublished manuscript by him held in the library and archives of London's Natural History Museum. Moreover, this beautiful painting remained unpublished until as recently as 2010, when it appeared in a Zeitschrift des Kolner Zoos article on quaggas by Lothar Schwahle and Wolfgang Wozniak.

Hamilton Smith's two illustrations readily confirm my early deductions as to the isabella quagga's likely appearance – namely, an aberrantly pale, isabelline-coloured quagga with only very faint, white striping.

Having viewed several comprehensive lists of quagga material currently housed in museums worldwide, I can confirm Lydekker's statement that the isabella quagga skin deposited by Burchell at what is now London's Natural History Museum is no longer there, and is therefore lost. Presumably it was discarded due to its very poor condition, but a tragic loss nonetheless of such an exceptional, unique specimen, and which nowadays might well have yielded much useful information via DNA tests conducte3d upon samples of this skin's tissues.

Yet despite the isabella quagga having long since been reduced in status from a taxonomically-discrete species to a non-taxonomic mutant oddity, its delicate pallid beauty deserves to be remembered and celebrated. So I am very glad that I discovered this elegant animal hidden away as the briefest of footnotes within the dusty archives of the past, and have been able to revive it, even if only in words and pictures, within this present article, written up at last.

 

Alongside a mounted quagga specimen at Tring Natural History Museum, England (© Dr Karl Shuker)

 

LET'S ALL DO THE KUNGA!

 

Detail from 'War' panel of Standard of Ur mosaic, c2600 BCE, depicting kungas pulling 4-wheeled battle-wagon (© Zunkir/Wikipedia – CC BY-SA 4.0)

It's always good to learn of a 'new' animal, even if it turns out to be one that actually existed four and a half millennia ago! And so, with no further ado, please welcome to ShukerNature the kunga.

Depicted in Mesopotamian art and referred to in cuneiform writings dating back 4500 years from the Fertile Crescent region of the Middle East, the long-vanished kunga was a powerful horse-like creature that was used as a draft animal to pull war wagons into battle and royal chariots during ceremonial parades. However, it has long been a puzzle to archaeologists and zoologists alike.

 

Close-up of a kunga, as depicted in a detail from 'War' panel of Standard of Ur mosaic, dating from c2600 BCE (© Agricolae/Wikipedia – CC BY-SA 3.0)

This is because the domestic horse Equus caballus was not introduced into this region until 4000 years ago, and the only known equine beasts that did exist there at the time of the kunga were the domestic donkey E. africanus asinus and the Syrian wild ass (aka the hemippe) E. hemionus hemippus, both of which were smaller and far less sturdy than the kunga. So what was it?

 

A beautiful vintage colour painting by French artist Georges Severeyns of a Syrian wild ass (the smallest type of modern-day wild equid known to science), in Nouvelles Archives du Muséum d'histoire Naturelle, 1869 (public domain)

Unlike many mystery beasts, there are actually physical, tangible remains of the kunga in existence, because as it was such a valuable, useful work beast, specimens were sometimes buried alongside persons of high social status, with several kunga skeletons having been discovered at the northern Syrian burial complex of Umm el-Marra. Unfortunately, however, the bones of kungas, donkeys, wild asses, and horses as well as mules and other equine hybrids, all look very similar, so although kunga bones have been examined, no conclusive identification of what the kunga actually was has ever resulted – until earlier this year.

In January 2022, published research in the journal Science Advances revealed that DNA samples had been extracted from the bones of one buried kunga specimen from Umm el-Marra and subjected to comparative sequencing analyses with other equine forms. These analyses confirmed that it was a hybrid – most probably the offspring of an interspecific mating between a male Syrian wild ass and a female domestic donkey.

 

Vintage engraving from 1841 depicting some Syrian wild asses in the wild state (public domain)

Moreover, because such matings would not occur naturally, and as the kunga, being a hybrid, was almost certainly sterile (just like the mule), male wild asses would need to have been deliberately captured and mated with female domestic donkeys on a regular basis in order to perpetuate the kunga strain 4500 years ago in the Fertile Crescent – thereby making the kunga the earliest recorded human-engineered hybrid. But when the horse was introduced here 500 years later, the kunga was no longer needed, so the breeding of it ceased, and this remarkable creature duly vanished from existence.

 

Syrian wild ass being captured by Assyrians, 7th Century BCE art found at Nineveh (Wikipedia/no restrictions)

Of course, one could ask why, now that the kunga's precise hybrid nature is finally known, scientists are not excitedly proclaiming "Let's all do the kunga – let's resurrect it!", by once again mating male Syrian wild asses with female domestic donkeys. The answer is as tragic as it is simple – no such restoration can take place because the kunga's paternal progenitor, the Syrian wild ass, is itself now extinct, as a result of over-hunting, with the last two known specimens both dying in 1927.

So unless the kunga and/or the Syrian wild ass can be recreated via genetic means in the laboratory one day, the kunga is doomed forever to remain nothing more vital than a collection of long-buried bones, a few brief mentions in cuneiform script, and some preserved images in ancient art. RIP.

 

A trio of poignant photographs depicting one of the world's last known Syrian wild asses, a specimen living at Vienna's Tiergarten Schönbrunn in 1915, plus one photograph (at bottom of photo-column) of a Syrian wild ass at London Zoo in 1872 (public domain)

 

 

THE ZEBRO - AN EQUINE MYSTERY FROM IBERIA

Artistic representation of the zebro's possible appearance ((c) ejbeneite.blogspot.co.uk)

During the Middle Ages and the Renaissance, several Spanish hunting treatises alluded to a mysterious, now-vanished equine creature known as the zebro (or encebro, in Aragon), living wild in the Iberian Peninsula. In one of these works, it was described as “an animal resembling a mare, of grey colour with a black band running along the spine and a dark muzzle". Others likened it to a donkey but louder, stronger, and much faster, with a notable temper, and whose hair was streaked with grey and white on its back and legs. What could it have been?

Although largely forgotten nowadays, the zebro experienced a brief revival of interest from science in 1992. That was when archaeologists Carlos Nores and Corina von Lettow-Vorbeck Liesau published a very thought-provoking article in the Spanish scientific magazine Archaeofauna, in which they boldly proposed that the zebro may have been one and the same as an equally enigmatic fossil species – Equus hydruntinus, the European wild ass.

Artistic representation of the European wild ass Equus hydruntinus (Francesco Gabelione, (c) 2009, IBAM ITLab)

The precise taxonomic affinities of this latter equid have yet to be satisfactorily resolved, for although genetic and morphological analyses suggest that it was very closely related to the onager E. hemionus, one of several species of Asiatic wild ass, it can apparently be differentiated from these and also from African wild asses by way of its distinctive molars and its relatively short nares (nasal passages). Arising during the mid-Pleistocene epoch, approximately 300,000 years Before Present, the European wild ass persisted into the early Holocene before finally becoming extinct. During the late Pleistocene, its zoogeographical distribution in western Eurasia stretched from Iran in the Middle East into much of Europe, reaching as far north as Germany, and it was particularly abundant along the Mediterranean, with fossil remains having been recovered from Turkey, Sicily, Spain, Portugal, and France.

According to Nores and Liesau, moreover, this species may have survived in southernmost Spain and certain remote parts of Portugal until as late as the 16th Century (they consider its disappearance to represent the Iberian Peninsula’s last megafaunal extinction), where, they suggest, it became known locally as the zebro. More recently, their theory gained support from the discovery of E. hydruntinus remains at Cerro de la Virgen, Granada, dating from as late as the 9th Century.

A sorraia stallion (Selona/Wikipedia)

Some researchers have also suggested that before dying out, the zebro gave rise at least in part to a primitive, nowadays-endangered Iberian breed of donkey-like domestic horse called the sorraia (which was once itself referred to as the zebro). Furthermore, many believe that it was from the term ‘zebro’ that ‘zebra’ originated as the almost universally-used common name for Africa’s familiar striped equids.

Even today, many Iberian place-names still exist in which the mysterious but now-obscure zebro’s name is preserved. These include Ribeira do Zebro in Portugal; and Valdencebro (in Teruel), Cebreros (Ávila), Encebras (Alicante), and Las Encebras (Murcia) in Spain.

A partbred grullo Sorraia colt exhibiting zebra type stripes formed by the lay pattern of the newborn foal hair coat. The dorsal and leg stripes remain, but the lay pattern stripes disappears after several days or weeks when the hair is fully fluffed (Selona/Wikipedia)

FURTHER READING:

NORES, Carlos & LIESAU, Corina (1992). La zoologia historica como complemento de la arqueozoologia. El caso del zebro. Archaeofauna, vol. 1, pp. 61-71.

This ShukerNature blog post was originally published as an article in issue #1 of Flying Snake.