Saturday, 6 July 2019

Taxon Surrogate Profiles North America # 1: Equidae


            Hey everyone. I’ve hit a bit of a writer’s block with the European series, so I’ve decided to start the next one and come back to it later. The next five articles are going to be explorations of the rewilding potential for five taxonomic families that have been completely (or partially in one case) extirpated from temperate North America. These are, in order of discussion, the Equidae, Camelidae, Tayassuidae, Tapiridae, and Elephantidae. For today’s article we will be starting with the Equidae, a family which owes its origins and much of its evolutionary history to North America, and yet is no longer found there in a truly wild state, having been extirpated sometime during or after the Pleistocene-Holocene transition. I will discuss the taxonomic identity of the mid-late Pleistocene  equids of North America, and the reintroduction possibilities for this group.
            The taxonomic history of the North American horses has been long and complex. It has been repeatedly confused by the presence of feral European animals, repeated prehistoric migration between North America and Eurasia, and the tendency of palaeontologists to radically overclassify. In the past few decades. We’ve gone from having dozens of recognized native North American equids, with taxonomic affinities variously ascribed to horses, asses, and zebras, to having only two legitimate species, one of which is still very much with us. Basically all fossil equids from the mid-late Pleistocene of North America can be placed into two groups. The first is the caballine group, which is now known to be of the same species as the living Equus caballus, which includes many extant and extinct groups of animals from Eurasia, including the modern domestic horse (which descends from the extinct European wild horse or tarpan) and the wild Przewalski’s horse, or takhi, from Asia. The second group is the stilt-legged equids, which were originally thought to be members of the living Asinus (sub)genus, which includes the Eurasian and African wild asses (Asinus hemionus and Asinus asinus), as well as the domestic African ass, or donkey. This conclusion was arrived at through morphological analysis, but genetic studies have since shown that this group is distinct and deserves its own genus. Consequently all stilt-legged equids from the mid-late Pleistocene of North America are referred to as variations of the single species Harringtonhippus francisci. With this information we can now refer any equid remains from this time period to one of these two groups.
            The caballines have been, in a sense, already reintroduced. Areas of North America once occupied by various extinct subspecies or varieties of Equus caballus (ex. complicatus, conversidens, niobrarensis, lambei, occidentalis, scotti, etc) were very quickly recolonized by feral Spanish horses, Equus caballus caballus, after said horses were brought to North America by the conquistadors. These populations resulted in the modern mustang, which was quite happily able to recolonize its former range through a combination of natural migrations and trading by First Nations peoples. These days however, mustangs are quite rare, and have been severely reduced in number and frequently crossed with other breeds. Some herds have purposely had individuals with certain colourations removed in order to create a more “primitive” appearance in the remaining herds, leading some to assert that the animals have always looked this way and that their primitive markings are the result of natural selection, when in reality the breed remains quite domestic and can easily be brought back into captivity. Knowing this, and considering their reduced population, I think other options could be explored when selecting horses for an experimental rewilding site. Considering their truly wild status (which has been unsuccessfully challenged in recent years), their truly primitive appearance and behaviour, and their adaptations to (semi-)arid steppes and semi-deserts, which represent much of the available habitat in central North America, I believe the Asiatic wild horse (Equus caballus przewalski) would be an excellent choice for introduction to North American rewilding areas. Having made a considerable recovery from their near-extinction in the first half of the twentieth century, there exists a sizeable captive population in North America which could be used to establish a free-living breeding population on the Great Plains or adjacent habitats.
This would run contrary to my suggestions for rewilding domestic horses or crossbreeds in Europe. This is for two reasons: the first being that domestic horses are more closely related to the European wild horse (Equus caballus ferus) than is the Asiatic wild horse, and the second being that most of the available habitat in Europe is different from what takhis are adapted to, and more easily occupied by native horses which have developed appropriate ecotypes. Genetic evidence suggests that the two living horse subspecies are about equally related to Equus caballus lambei/lenensis, which occupied the Bering strait, and are consequently likely to be equally related to the other North American horse subspecies, since Beringia would have represented the migration corridor between Eurasia and North America. Knowing this, it seems more appropriate to select the taxon that is already considered wild, could use the conservation support, and will not require any significant selection. The fact that takhis already look like primitive wild animals will also be useful for rewilding projects funded by ecotourism. Their ability to occupy non-steppic habitats is established from their introduction to conservation areas in Europe with varying levels of forest cover. Probably the best course of action is to begin with the central plains, where most American rewilding projects are focused/planned anyway, and work outwards from there. Their ability to thrive in colder climates, such as those that might be experienced in rewilding areas in the far north, will need to be assessed. However, considering we now have the whole genome of an extinct relative adapted to these areas, genetic hybridization might be an angle. Hybridization with existing domestic breeds adapted to these areas, such as the Yakut pony, would certainly be easier, but it would also potentially introduce some less desirable behavioural traits. The best course of action will be decided when relevant.
As to the replacement for Harringtonhippus, the substitute used will be a generic proxy, rather than a subspecific one. Based on morphological and chromosomal differences, I think it appropriate that the various Equus subgenera be elevated to genus level, including Asinus, the group that includes the wild asses. Although not actually a member of this group, the North American stilt legged equids shared some remarkable morphological similarities with Old-World stilt-legged horses surviving today, and seem to have occupied similar open and arid habitats based on fossil records. Knowing this, I think the best proxies would be various subspecies of Asinus hemionus, the Eurasian wild ass, which were once widespread across multiple habitat types in Europe and Asia. While the kulan and onager subspecies have been proposed for redistribution in Europe and western Asia, and the khur subspecies is in the process of being reintroduced to new sites in India, the Mongolian hemionus subspecies and the Tibetan kiang subspecies might potentially be used in North American rewilding projects, with the former being well suited to the prairies and deserts of northern Mexico, the central/western United States, and southern Canada, and the latter being adapted to colder habitats such as those found in northern Canada and Alaska. These two subspecies are the most populous of any of the Eurasian wild ass varieties, with almost 70,000 of the kiang subspecies remaining in the wild, and almost 20,000 of the hemionus subspecies. Oddly enough however, they are also the least studied taxa, and the rarest in captivity. Consequently, individuals used in rewilding projects in North America would probably have to be wild-caught, which would be advantageous for acclimatization purposes, but expensive due to the capture and transportation costs.
There are many potential benefits to equid reintroduction in North America. Horses are an important part of the grazing regime in any ecosystem where they occur. Their ability to eat tougher and poorer quality forage allows them to open up habitats for more selective herbivores. Feral mustangs were often followed by herds of bison for this exact reason, in a dynamic mirroring that of zebras and wildebeest in Africa. Something worth considering however is that predation by wolves and pumas would likely be insufficient for population control and human intervention would possibly be necessary to keep numbers in check. The main predator of the horse in the Pleistocene was the American lion (Panthera (leo) atrox). North America probably has a better chance of returning lions to its ecosystems, at least in fenced game reserves, than does Europe, and I for one would be very excited to see that particular predator-prey relationship first-hand. A truly rewilded Great Plains could potentially become one of the greatest ecotourism destinations on Earth.

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