Friday, 12 July 2019

Taxon Surrogate Profiles North America # 2: Camelidae

         One of only three families of Artiodactyls to have evolved in North America (the others being the Antilocapridae and Tayassuidae), the Camelidae are amongst the most distinctive members of the even-toed ungulates, being more distantly removed from cattle, deer, and pigs than even whales are. Despite having originated in North America, and being found there exclusively for roughly ninety percent of their existence, the Camelidae no longer have native representatives on the continent, and haven’t for over ten thousand years. During the late Pleistocene, both living tribes of Camelids, the Camelini and Lamini, were found in North America, with four to six genera split between the two groups. The camels were represented by Camelops, Paracamelus, and Titanotylopus (possibly distinct from Gigantocamelus, unclear), and the llamas by Hemiaucheniaand Palaeolama
Camelops hesternus, known colloquially as the western camel, would have been roughly the same size as members of the living genus Camelus, and was a typical inhabitant of dry, open habitats in western North America up until the Pleistocene-Holocene transition. Paracamelus, the genus to which the modern species owe their ancestry, could still be found in the more northern areas of the continent, in the form of Paracamelus gigas, during the middle Pleistocene, though their exact date of extinction remains unclear. Paracemelus crossed the Bering land bridge around six million years ago, at which point it quickly spread across the arid and semi-arid regions of the Old World and eventually evolved into the modern Camelus, the genus to which living Dromedary and Bactrian camels belong. Camelops also ranged into these areas during interglacials, but was not present during the colder stages and consequently it never crossed the Bering strait into Eurasia. Titanotylopus nebraskensis was the largest of the Pleistocene camels, standing three and a half meters high at the shoulder, and weighing almost three metric tonnes. The official date of extinction for this group is sometime in the Irvingtonian period, but some recent finds from Alberta may push their occurrence well into the Rancholabrean. All three of these Cameline genera are thought to have been open-country browsers, much like the living genus. Hemiauchenia contained many species very similar to the llamas and alpacas (plus their wild counterparts) that are still with us today, and in fact these animals are the descendants of members of this genus which colonized South America during the Great American Interchange roughly five million years ago. In the late Pleistocene one species, Hemiauchenia macrocephala, which would have resembled a wild llama but about thirty percent larger, was still quite common across western and southern North America. Palaeolama was also a member of the Lamini tribe, but unlike Hemiauchenia or its descendants, which are/were largely mixed feeders adapted to open environments, Palaeolama appears to have been a strict browser adapted to closed or semi-closed forest environments, a niche not filled by any living camelid today. The only species of this genus that made it to the Pleistocene-Holocene transition in North America was Palaeolama mirifica, which was found in the American southeast. Members of both Hemiauchenia and Palaeolama also existed in South America during this period, as did a poorly described genus known as Eulamaops, an alpaca-sized camelid that appears to have been an open-country browser.
Ever since Pleistocene Rewilding first arose as a concept, camels (alongside horses, lions, cheetahs, and elephants), have been touted as a no-brainer for reintroduction to the American west. Generally, most people who subscribe to the idea of trophic rewilding using pre-Holocene baselines believe that camelids belong in North American ecosystems, but correct implementation of said introduction evades common consensus. Some say that Dromedaries (Camelus dromedarius), with their ease of availability, are the best choice. Others say domestic Bactrians (Camelus bactrianus), which are also available commercially (albeit less cheaply) and which originate from very similar habitats and climates to those found in western North America, would be preferable. My belief is that the best options for a combined Camelops-Paracamelus-Titanotylopus substitute is the last wild member of the Camelini, the wild Bactrian camel, Camelus (bactrianus?) ferus, and there are several reasons for this. The first is that the wild Bactrian is heavily threatened in its native habit in China and Mongolia, being considered Critically Endangered by the IUCN, with just over a thousand individuals remaining, split between two or three populations, and only one captive breeding program. The creation of a fourth population on the Great Plains, which is closer to what is generally considered ideal camel habitat (as opposed to some of the extreme desert habitats where human activity has pushed them) and where poaching and hybridization with domestics would largely be a non-issue, could potentially be very beneficial to the future of the species.
The second reason why wild Bactrians might be preferable is that, as a truly wild and non-domestic species, they are probably better prepared for wild-living in a landscape with significant predator densities. It is true that domestic camels go feral quite easily, but their interaction with large predators is not well-documented, with the most famous feral camel population, the Australian Dromedaries, not having to deal with anything larger than a dingo, which is completely ineffectual at controlling camel numbers. Camels in East Africa occasionally have conflicts with lions, but these populations are rarely considered feral, and individuals are generally quite helpless in their behaviour. Wild and domestic Bactrians may deal with wolves occasionally, but these mostly target calves, with the same being true for bears. In the distant past (before the Holocene), wild camels would have had to deal with cave lions (Panthera (leo?) spelaea) and hyenas (Crocuta crocuta spelaea), and in historic times when wild camels still roamed the Caspian region they would have occasionally experienced predation by tigers and leopards. The camel’s size and aggressiveness are its main defences against predators, which will usually target smaller prey when available. 
Obviously obtaining wild Bactrians would be difficult (though starting a captive population really should be a priority regardless) but I think it’s alright to play the long-game in this case. Projects in Mongolia have shown that they can be kept and bred in captivity which, regardless of rewilding potential, would possibly represent an opportunity to begin replacing domestic Bactrians and Dromedaries (both of which have little conservation value) in accredited zoological collections with wild Bactrians. A similar process is ongoing with the replacement of captive collared peccaries (Pecari tajacu) with Chacoan peccaries (Catagonus wagneri), due to the Endangered status of the latter species, compared to the Least Concern status of the former species. Breeding the wild Bactrian in Canada, Mexico, and the US would be a great first step for a potential introduction to wild areas in the North American continent, as well as a potentially crucial step in the preservation of the species.
Reintroducing the Lamines to North America is considerably less complicated. Wild llamas (guanacos, Lama guanicoe) and wild alpacas (vicunas, Vicugna vicugna) are both considered Least Concern. Consequently, even though their domestic counterparts would still be easier to obtain, it makes far more sense to acquire wild animals, as it would be completely feasible even in the short-term. There exists already a large population of guanacos both in zoos and in private collections in North America, and a wild population could conceivably be sourced from these institutions. Vicunas do exist in captivity in North America, but in very low numbers, and more would probably have to be sourced either from the wild or from zoological collections in Europe, where the species is more numerous. Both are native to similar habitats to those found in western/southern North America, and guanacos also inhabited similar climates to that of the south-eastern US in prehistoric times. Both species are adapted to predation by large cats, such as pumas and in some places jaguars, and to a lesser extent also bears and large canids (of which there were more species in the past).
I’m kind of abandoning the idea of direct proxies in this case. Camels are being used as a combined proxy for multiple genera that have occupied the continent at various times, with the understanding that its ecological role will be similar but perhaps not directly analogous to these groups. Both guanacos and vicunas are the descendants of Hemiauchenia, but are smaller and, especially in the case of the vicuna, probably differing in their exact dietary choices. The idea here is more about returning the family and its remaining wild representatives to a continent where they were previously prevalent and ecologically important, with the hope that they will create their own unique dynamic with the ecosystem that is similar but not identical to that displayed by their extinct relatives. 
That being said, Camelus ferus and Lama guanicoe would probably be functionally and aesthetically very similar to Camelops hesternus and Hemiauchenia macrocephala respectively. The western camel is often depicted in palaeoart as Dromedary-like in its appearance, but this is likely an incorrect representation. Dromedaries are very derived and have adapted for hot, tropical deserts, an ecosystem not familiar to its ancestors, which were inhabitants mostly of temperate steppes and semi-deserts, such as those occupied by Bactrians. Consequently western camels may have had similar adaptations to these habitats, such as long winter coats. Dromedaries also have only one hump, which has been shown to be a derived trait, with foetal Dromedaries developing and then losing their second hump in the womb, and adult males occasionally displaying a vestigial hump above their shoulders. This has been taken as evidence that one-humped camels evolved from two-humped ones, and that having two humps is consequently the more primitive state. This would mean that if North American Camelines had humps at all, and their spines suggest they did, it would be far more parsimonious for them to have two humps, rather than one. Large-headed llamas (Hemiauchenia macrocephalafor their part, probably looked a lot like tall guanacos, with slightly different skull and leg proportions.
Camelids offer a unique ecological service not common amongst the remaining large mammals. Their extreme lack of selectiveness when browsing allows them to be effective controls and distributors of numerous plant species that other herbivores won’t even touch. Dromedaries employed by the US army in the American southwest during the nineteenth century were observed to eat creosote (Larrea) and mesquite (Prosopis), and camels in this same area are sometimes used to control these species along with cactus (Opuntia), saltbush (Atriplex), and others. Camels and llamas will eat the thorniest, driest, toughest plants around, opening up space for other plants more useful to other herbivores and creating more mosaic ecosystems. After these species are introduced, we might begin see a dynamic develop between Camelids, Equids, and ruminants, where camels and llamas enter overgrown areas first to clear out all the scrubs and brush, and are then followed by horses and onagers to eat all of the taller, tougher grasses that pop up, to then be followed by bison and elk to eat the shorter, more nutritious grasses that grow up when all the other plants have been eaten and trampled. This dynamic would become more complex with the addition of other native and introduced herbivores, and is of course a simplification of how the real process might work, but would nevertheless potentially create some very high biodiversity in comparatively small areas.
There are also just the generic benefits that could possibly be obtained from the introduction of any sort of megafauna to the continent, namely ecotourism, sustainable offtake, and all of the numerous ecological and economic benefits I’ve covered in previous articles.  As large and charismatic animals, with unique appearances and behaviours, camels and llamas should be a huge draw for ecotourism in a rewilded American prairie. Much like the Equids discussed in the last article, camelids would represent the return of a taxon which owes its origins and most of its evolutionary history to the North American continent. I subscribe to the theory that their extinction, and that of numerous others, was ultimately anthropogenic, and it is my belief that filling some of these vacant niches through translocation is one of our greatest potential strategies against the ongoing loss of global biodiversity. North America may potentially have one of the highest diversities of large herbivores in the world someday, with representatives from as many as eight different families having an ecological claim to the continent.

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.

Tuesday, 18 June 2019

Taxon Surrogate Profiles Europe # 6: Caprinae Part 1

          The Caprinae are a subfamily of the Bovidae family which includes goats, sheep, and their relatives. In modern times there are three genera of the Caprinae that are considered native to Europe, including a combined seven species. The first is the Capragenus, which includes the four European species of ibex: the Anatolian ibex (Capra aegagrus), the Caucasian ibex (Capra caucasica), the Alpine ibex (Capra ibex), and the Iberian/Pyrenean ibex (Capra pyrenaica). The second genus is Rupicapra, which includes the two species of chamois: the Iberian/Pyrenean chamois (Rupicapra pyrenaica) and the European chamois (Rupicapra rupicapra). The last genus is Ovis, which includes the wild sheep, or mouflon (Ovis orientalis). The geographic history of these species within Europe is complicated, and there was a fourth genus present in the area during the mid-late Pleistocene, which will also be discussed. Muskoxen and their extinct relatives (Ovibos, Praeovibos, Soergelia, etc) are also part of this subfamily, but they will be discussed in the next article.
            First, the ibex. The genus Capra was once found across every mountain range and rocky habitat in central and southern Europe. The Anatolian subspecies of the bezoar ibex (Capra aegagrus aegagrus), the smallest of the European ibex species at a maximum weight of ~90 kg, is found in the rocky habitats of Turkey and adjacent Armenia, Azerbaijan, and Georgia, in addition to several countries not generally considered to be part of the European continent. This taxon represents the ancestral stock from which all domestic goats descend, and is still found in a large portion of its previous distribution. Despite this, it is considered vulnerable by the IUCN and has suffered population reductions in some regions due to poaching, trophy hunting, habitat loss, and hybridization with its domestic relatives. These factors have led to its complete extirpation from several areas, including most of western and northern Turkey. Nearby, in the Caucasus mountains, including Georgia and the adjacent regions of Russia, the Caucasian ibex is found. Traditionally this species was split into two, one for the western part of the mountain range and one for the east, but this has since been reconsidered so that that they are now considered subspecies: the western Caucasian ibex (Capra caucasica caucasica) and the eastern Caucasian ibex (Capra caucasica cylindricornis). At a maximum weight of ~140 kg, this is the largest of the European ibex species. While the western subspecies is considered endangered, the eastern is listed only as near-threatened. This species has the smallest original distribution of the four European ibex species, and consequently has the fewest opportunities for reintroduction. The protection of its existing habitat is very important to it continued survival.
            The Alpine ibex, of which there are no recognized subspecies and which is considered to be least concern by the IUCN, seems to have suffered the greatest range reduction out of these four species. Present only in the Alps today, this species was also present in the Tatras, Balkans, and Dinaric mountain ranges until the mid-Holocene, and the Carpathian and Apennine mountain ranges until at least the late Pleistocene. Extirpated from everywhere but the French and Italian Alps by the early twentieth century, the ibex has since been reintroduced to the alpine regions of Austria, Germany, and Slovenia. Reintroduction attempts have also been made in Slovakia to replace the extinct Tatras population, but the project failed due to the animals being not only of Alpine stock, but also of Anatolian and even Nubian (Capra nubiana) individuals. At the time these ibex varieties were considered to be subspecies of each other, and it was thought that a broad genetic base would increase the likelihood of a successful introduction. However, the hybrids molted at the wrong time of the year, due to the adaptations inherited from their more desert-adapted ancestors, and the population did not survive the winter. Future attempts should use only non-hybridized C. ibex. A population also exists in Bulgaria, imported for sport. These animals are often treated as alien, but their Holocene fossil record in the area suggests that they are better thought of as a reintroduced species. Further reintroductions to the Balkans and Dinarides should be a part of any rewilding projects in those areas. I would also put forward that the Apennines and Carpathians should be considered for reintroductions as well.
            The last of the European ibex species is the Iberian ibex. In modern times there have been four subspecies of this ibex living in the Iberian peninsula. These were the eastern Spanish ibex (Capra pyrenaica hispanica), the Portuguese ibex (Capra pyrenaica lusitanica), the Pyrenean ibex (Capra pyrenaica pyrenaica), and the western Spanish ibex (Capra pyrenaica victoriae). Of these, the eastern variety is considered least concern, the western is vulnerable, and the Portuguese and Pyrenean are extinct, the former since the late nineteenth century, and the latter since the early twenty-first century. Both were victims of hunting, competition with livestock, and habitat loss. The Pyrenean subspecies is notable for having gone extinct twice: first in 2000 when the last wild individual was hit by a fallen tree, and again in 2003 when a kid was cloned from material taken from this individual, only to die after several minutes. It is my opinion that future cloning efforts should be reserved for species with more unique evolutionary/ecological histories, and that the best option for the Pyrenean ibex is to create new populations using another subspecies, something that has already been considered by French organizations. It may be possible to integrate the genetic material of the Pyrenean subspecies into these populations in the future. The Portuguese subspecies is in the process of being replaced by the western ibex subspecies through natural dispersal and translocation.
            Moving on to the chamois group. The Pyrenean chamoi has three living subspecies, all of which are considered least concern by the IUCN, these are: the nominate subspecies (Rupicapra pyrenaica pyrenaica), which occupies the Pyrenean mountains of southwestern France and north-eastern Spain, the Cantabrian chamois of north-western Spain (Rupicapra pyrenaica parva), and the Abruzzo chamois (Rupicapra pyrenaica ornata) of the Italian Apennines. Fossil records indicate that the Pyrenean chamois used to occupy the entirety of the Iberian peninsula until they were extirpated from Portugal and most of Spain in the middle Holocene. It is unclear if these populations were referable to the Cantabrian subspecies or to some extinct group, but either way the possibility for a redistribution of the species across the peninsula could be possible, especially with the increasingly low population densities in some areas of Iberia. The other chamois species has a much larger range and a greater diversity of subspecies. The nominate subspecies (Rupicapra rupicapra rupicapra) is found in the Alps, and there are also Balkan (Rupicapra rupicapra balcanica), Carpathian (Rupicapra rupicapra carpatica), Carthusian (Rupicapra rupicapra cartusiana), Anatolian (Rupicapra rupicapra asiatica), and Caucasian subspecies (Rupicapra rupicapra caucasica), in addition to a critically-endangered subspecies from the Tatras (Rupicapra rupicapra tatrica). Populations of all these taxa have undergone reductions in their distributions, and have become heavily fragmented by habitat loss. A priority for their recovery will be to improve connectivity between their pockets of occurrence.
            As for the third genus, Ovis, the history in Europe is a little more complicated. The current representative of the wild sheep in Europe is the European mouflon (Ovis orientalis musimon). This population descends from wild sheep found on the islands of Corsica and Sardinia, which were later imported to the mainland for hunting purposes, where they have since become very widespread. The problem is that the original stock were not as wild as originally thought, being the descendants of an early variety of domestic sheep (Ovis orientalis aries), which were able to go feral on these islands only due to the lack of large predators. They display very domestic behaviours, preferring lowland habitats to rocky highlands, and being very easily taken by predators, so much so that when predators have returned to certain areas in the past, the local mouflon population has disappeared completely. This needs to be considered if mouflon are to be used in rewilding projects. One approach is to use one of the truly wild subspecies instead, such as the wild sheep of the Transcaucasian region (Ovis orientalis gmelini) which should be better adapted to predation and to harsh terrain. This subspecies is threatened, and expanding its distribution would certainly help in its conservation. We would have to be prepared for the possibility of hybridization with existing populations of the European variety, but this is to be expected, and it is possible that a combination of the two varieties would be quite suitable for multiple European regions, assuming that natural selection will favour crossbreeds which inherit wildtype behaviours. The original distribution of mouflon on the mainland included the Balkan and Italian peninsula, in addition to Turkey and Transcaucasia, but fossils of the genus Ovis have been found outside of this region as well, allowing for the possibility of their use in rewilding projects in most of the European mountain ranges. 
            The last genus I will be discussing in this article is one that no longer occurs in Europe, and that is Hemitragus, today represented only by the Himalayan tahr (Hemitragus jemlahicus). From the mid-late Pleistocene there was another species in this genus, known as the European tahr (Hemitragus bonali). This species would have coexisted with the other three genera, and had a range that included all of the same mountain ranges. There even exists cave art that appears to depict a tahr (alongside an ibex), displaying many of the features that define the living species, such as short horns and a thick mane of fur around the neck. Native only to the Himalayas today, the tahr has become an invasive species in many areas where it has been imported for game hunting, including Argentina, New Zealand, South Africa, and the southern United States. This ability to adapt to a range of habitats speaks well for their potential introduction to Europe as a replacement for their extinct relatives. It is unknown exactly how they would interact with the other Caprines, but its relative did so seemingly without competitive exclusion for hundreds of millenia, and the Himalayan species is sympatric with several related species, such as (depending on the region) bharal (Pseudois nayaur), argali (Ovis ammon), goral (Nemorhaedus goral), serow (Capricornis thar), and markhor (Capra falconeri). The ibex, mouflon, and chamois seem to avoid competition through living at different altitudes and through differences in the amount of browse or graze taken by each species. Presumably the European tahr would have fit into this dynamic somehow, but more research will be necessary to determine how, and if the living species would have similar interactions with its environment.
            Populations of all four genera would be predated on by wolves (Canis lupus), bears (Ursus arctos), and lynx (Lynx lynx). Their re-establishment would also be an important preliminary step in the reintroduction of the leopard (Panthera pardus) to European rewilding areas. Human hunting of goat-like species is already a huge industry, and expansions and introductions would only reinvigorate that industry, with the understanding that offtake has to be sustainable. Wild meat and leather could be sourced from the offtake as well. These are also all species that people would potentially enjoy seeing in natural areas, and which could further stimulate ecotourism. Their ability to traverse uneven terrain will make them especially important for fighting succession and stimulating nutrient cycling in high altitude regions where larger herbivores like red deer and bison are less able to traverse the terrain. 

Friday, 14 June 2019

Taxon Surrogate Profiles Europe # 5: Bubalus

            Now for the last of the European bovines, the water buffalo. Unlike the bison (Bos bison) or aurochs (Bos taurus), the European water buffalo (Bubalus (bubalis?) murrensis) never returned to Europe after the last interglacial and eventually went extinct, possibly due to human presence in their glacial refugia. However the wild Asian water buffalo (Bubalus bubalis cf. arnee) made it to the eastern regions of Turkey, the Levant, and possibly the southern Caucasus during the early-mid Holocene, and the domestic descendants of the Asian species (Bubalis bubalis bubalis) have been common in eastern and south-eastern Europe for over a thousand years, where they have developed into robust and cold-resistant landraces. It would appear that buffalo still have an open niche in the European landscape, being more inclined to living in wetland habitats and consuming aquatic plants than their relatives in the genus Bos. It is unknown whether the extinction of the European species, which had disappeared from Europe and reappeared later several times as interglacial stages came and went, was ultimately due to human hunting or whether the species was naturally replaced in its refuges by the modern species, which was then prevented from recolonizing Europe by human activity. Frankly I’m not sure it matters, since the niche remains unfilled and either way it seems to me that humans are at least partially responsible for the lack of water buffaloes in Europe today.
            The question would then be how to go about restoring the water buffalo niche to the continent. The process is already partially under way in the form of local domestic buffalo being used in grazing projects for wetlands in the Netherlands, Romania, and elsewhere. These animals are resistant to European pathology, and acclimated to its temperatures and available forage. Much like horses, and unlike cattle, they retain a less derived, less heterogenous appearance, and distinguishing them from wild animals would be difficult to the untrained observer. Consequently, converting these herds to an aesthetically and behaviourally appropriate wild population is considerably less difficult. Feral water buffalo are already quite common in many places around the world, but seeing as their wild progenitors are still very much with us, we have the option to explore hybridization as a means of expediting the process. The purpose of hybridization in this case is mostly just to increase the size of the animals, which have undergone the standard dwarfing associated with domestication, and to reintroduce wildtype behavioural genes. Now, the European breeds of water buffalo are all of the river type (Bubalus bubalis bubalis), as opposed to the swamp type (Bubalus bubalis carabao). This is because they descend from animals of Indian stock, which were domesticated from the Indian/Nepalese subspecies of wild buffalo, Bubalus bubalis arnee. The swamp type buffalo has a different number of chromosomes, and was domesticated separately, probably from the Indo-Malayan subspecies, Bubalus bubalis theerapati. As a consequence, animals used for hybridization should be of the arnee type, which I believe are the only variety present in European zoos in any case. These will have the same number of chromosomes as the domestic stock we’ll be using, and will also be adapted to slightly cooler habitats, such as those they might encounter in the Himalayan foothills. Due to the fact that the introgression would be coming from the ancestor taxon, rather than just a related taxon in this case, it is less important that the breeding process exclude sex chromosomes or mitochondrial DNA from the wild species, since they will likely be very similar anyway. Selection of the resulting crossbreeds will also be much less extensive, since all we want is a larger animal with uniform colour and non-domestic behaviour. Once the correct genetic material is there to build off of, natural selection will do most of the remaining work.
            The European water buffalo’s former distribution suggests that it used riverways, such as the Danube and the Rhine, as a means of dispersal. It was never able to reach the Italian or Iberian peninsulas, possibly due to the absence of suitable pathways and an inability to cross the Alps or the Pyrenees. Resettlement of the genus should begin in Central European river deltas (the Oder Delta or Danube Delta, for example), allowing for natural dispersal elsewhere. Their suitability for other areas where their relatives never occurred will have to be decided on a case-by-case basis, and should take into account that buffalo fossils are very rare in Europe, and can be confused with bison or ox fossils, obscuring their true distribution. Buffaloes have already proven incredibly useful in the management of wetland areas, and a self-sustaining wild population will allow these effects to be much more widespread and cost-efficient, as well as providing an additional species for invigorating tourism and sustainable hunting practices.

Saturday, 8 June 2019

Taxon Surrogate Profiles Europe # 4: Bos (taurus)

            Of all the proxies necessary to fully restore the European megafaunal assemblage, the aurochs (Bos taurus primigenius) is perhaps the most difficult to replicate in a satisfactory way. Attempts have been made for almost a century to breed them back in to existence through the crossing and selection of primitive cattle (Bos taurus taurus), but this has had limited success. Unlike horses, cattle do not go feral easily. Generations of selection for neoteny and docility have made even some of the least derived and demanding breeds feeble in comparison to their wild relatives. When they do live semi-feral they remain very habitat-specific and often require additional feeding. On top of that the traits known to have been possessed by the aurochs are spread very unevenly amongst its descendants, and animals showing more than half of them are very rare. You can certainly combine all of these traits in one population, but there is so much heterogeneity in cattle that getting them in one individual is almost impossible and getting them in enough animals to form a large founding herd has become a pipe dream. I’m going to posit some possible solutions, but I won’t pretend to know the best way.
            Probably the most straightforward way is to pursue cloning. We have DNA from several aurochs at this point and we can compare their genomes to that of living cattle. Unfortunately true cloning, wherein a living cell nucleus is implanted into a new egg and fertilized in a surrogate, is probably not possible as we would need an intact aurochs cell to start. Cloning only a few aurochs would be a poor start for genetic diversity as well, and you’d probably have to backcross them with cattle. However, direct splicing of wildtype alleles into a cattle genome might be possible if it were determined which genes were important. That might also be difficult considering how many genetic differences there might be, and the number of genes that might control things like body shape and horn size. There would be a lot of trial and error necessary to get it right, which is already the case with the breeding projects. Genetic engineering is also a very expensive process potentially, and though that may lessen over time, the effort necessary to identify important modifiers may remain high.
            Breeding primitive cattle over time is certainly cheaper, and perhaps more practical, but it definitely is not easy, and there are a million different opinions on the best way to do it. Forming a herd for conservation grazing using local cattle is easy enough, but these are still livestock under the law and will require additional feeding and veterinary care. Creating something that looks and behaves like a wild animal, and which can be left to its own devices with minimal human intervention, is considerably harder. There are certain breeds with a more aurochs-like phenotype than others, and this is especially true for some of the Lusitanian breeds like Sayaguesa (and their close relatives Serrana Negra and Alistana-Sanabresa), Tudanca, and Limia, but it also true for various breeds from across Europe. I certainly have my favourite breeds to use to meet this goal, but so does everyone else, and there will be issues regardless. Even the most primitive cattle have problems with hormonal expression, with this being less evident in fighting bull breeds, which still have some neotenous features. These lead to short faces, flat or concave backs with no shoulder hump, reduced sexual dichromatism, and long bodies with short legs. Basically, trying to breed an aurochs from a bull is liking trying to breed a wolf from a dog. You might get something that looks kind of like a wolf but it won’t act like one, and you’ll still get domestic features like floppy ears and white spots showing up.
            One potential method that might yield interesting results is hybridization. Now, I know that’s my solution for everything, but that’s only because I think it’s a legitimately useful tool for this kind of thing. Wild bovines have many of the traits we’re looking for in an aurochs effigy, there would just remain the question of which species to use. Bison (Bos bison ssp.) are out, since the aurochs and European bison were sympatric and we want to discourage hybridization in areas where both types of bovine are used. Yaks (Bos grunniensssp.) are in the bison lineage, so they should probably also be excluded. They do have some impressive horns, but they also possess many non-aurochs like traits associated with hair growth/colour and general body-form. Gaurs (Bos gaurus ssp.) share some ecological and morphological affinities with bison as well, and the kouprey (Bos sauveli) is likely extinct. That leaves the banteng (Bos javanicus). Now, while studies based on mitochondrial DNA place the banteng as being only distantly related to the aurochs, studies based on nuclear DNA, which I find more reliable, actually place it as being quite close phylogenetically. I think the Javan banteng (Bos javanicus javanicus), is the best variety to use, due to its perfect sexual dichromatism. The males are all dark brown to black, and the females all have an orangey-buff colour, a trait shared by the Eurasian aurochs. They are also the only banteng subspecies with the same number of chromosomes as cattle. An experiment breeding banteng to some of the more primitive Iberian breeds of cattle, like Sayaguesa or Maronesa, could potentially produce some very interesting results. There are two phenotypic traits that will have to be bred out, and potentially one or two behavioural ones, but the potential for hormonal improvement, non-domestic behaviour, and general robustness would make it worth it in my opinion. The white socks and rump of the banteng are dominant traits in existing cattle-banteng hybrids, making it an easy thing to breed out, which really just leaves the non-aurochs-like horns, which is already a trait that will need to be bred out in some of the primitive cattle breeds that might be used. Banteng also have a less confrontational style of fighting between bulls, but this may be selected out naturally. Something that is worth noting is that only the females of banteng-cattle crosses are fertile, and this will have to be accounted for in the breeding process. All backcrosses are fertile though, so that only affects the first generation hybrids. Apparently crosses with Indian cattle (Bos taurus indicus) which are also aurochs descendants, albeit of different subspecies (Bos taurus namadicus), produce fertile male offspring with banteng, but sources are a bit inconsistent on the matter. A hypothetical breeding plan would go as follows:

Male javanicusx Female taurus
-       F1 hybrid female, 1/2 javanicus, 1/2 taurus
-       javanicus X-Chromosome, taurus X-Chromosome and mDNA

Male taurusx Female F1
-       F2 hybrid male, 1/4javanicus, 3/4 taurus
-       taurus Y-Chromosome, either javanicusor taurus X-Chromosome, taurus mDNA
-       Select males that don’t show the banteng-type markings for breeding

Male F2 x Female taurus
-       F3 hybrid male, 1/8 javanicus, 7/8 taurus
-       taurus Y-Chromosome, taurus X-Chromosome and taurus mDNA
-       Integrate these animals into breeding-back herds to improve development and behaviour

Banteng are a tropical species, and this is worth noting for areas in central/northern Europe where cattle might be rewilded. Banteng are a hardy species, and cattle with slight banteng influence should do very well in most areas, as any climatic disadvantage should be counteracted by the adaptations of the base breed. For example, in the colder regions of the eastern steppe, a herd of 75% Podolian ancestry (ex. Maremmana, Podolica), 20% Lusitanian ancestry (ex. Sayaguesa, Limia), and 5% Javan ancestry might do very well. A herd in southern Spain might have similar proportions of Lusitanian and Javan ancestry, but be predominantly made of more arid-adapted Iberian breeds like Maronesa, Pajuna, or Lidia (fighting bull). What is important is that the herds can survive in the areas available to them without too much assistance, and that the desired traits are present in the herd to select upon over time. Natural selection will help this process along as well, as will occasional culling of bulls with an excess of undesired traits. I had originally planned to describe a similar plan to what I had proposed for horses, wherein different breeds become different varieties for the various European ecotypes, but due to the greater disparity in the appearance of wild vs domestic cattle when compared to horses, and the consequently smaller number of suitable breeds, there would probably end up being more like three or four types rather than nine, with herds in the colder regions of the north and east being descended from some of the same stock as those from founding herds in the south and west, only with greater proportions of ancestry from cold-adapted breeds. In general the strategy when forming new populations should, in my opinion, be to assemble herds of the most aurochs-like local cattle, or failing that the most aurochs-like cattle from a similar climate, and then slowly phase in breeds which possess desirable traits not found in the existing population. Certain breeds may be used to improve specific traits across several populations. As an example, the Chianina, from Italy, is one of the largest commercially-available breeds in the world and is itself a very old breed. Introgression from this breed, or from less common breeds of similar size (ex. Maltese cattle), could help to increase the size of aurochs-like cattle, which is important since the aurochs could be as tall as two metres at the shoulder, and weigh as much as a metric tonne, a size attained only by the largest of living cattle. Another breed, the Watusi of Africa, has also been used in breeding projects to contribute a greater horn size, as well as general robustness and improved ease of birth. The aurochs had very large horns and although many primitive European breeds possess horns of the right shape, they are of a smaller size which is a difficult thing to breed against if the genes aren’t there. The strategy with these breeds would not be to integrate them directly into existing herds, but to use them in the creation of individuals which will be integrated after a few generations of controlled breeding. 
            This is going to be a long process. It’s been going on, on and off, for almost a hundred years and we’re still not really there. Granted, we didn’t have the knowledge then that we have now, and progress has accelerated in the last twenty years, but cattle breed slowly and it can be very difficult to see progress in a short amount of time. It may be another fifty years of slow acclimatization and selection before we get an individual that’s sufficiently aurochs-like, and another fifty after that to get all of those traits in a population. We have to play the long game with this one, as is the case with many restoration efforts, but in the meantime we can build up primitive herds for use in our various European grazing projects and set the stage for the process to continue. This is going to be more difficult than it will be to re-establish herds of bison or water buffalo (which I will talk about in my next article), but it will be equally necessary for rewilding in areas where those species are not ideal. After all, all three varieties were once native to Europe, and each had their own unique ecological roles and habitat preferences. If you ask me, the more native megafauna we can restore, the better. Their environmental and economic benefits will become invaluable as land abandonment and vegetative succession continues.

Island Rewilding Series #1: Reunion Island

            The most common usage of purposeful taxon substitution in non-hypothetical situations thus far has been for the purpose of cons...