Return of the Wolf God Recently identified subspecies ranges farther than first imagined: across Africa. By Cheryl Lyn Dybas Stone compound in a traditional village in the Menz Guassa Community Conservation Area of the Ethiopian Highlands: The region is known for harboring the rare Ethiopian wolf, but genetic tests are revealing the presence in Ethiopia and elsewhere of a subspecies of gray wolf. Below: Canis lupus lupaster, the newly identified African wolf, was generally considered a subspecies of golden jackal until the recent discoveries. DNA testing of animals (which can be done on scat) is needed to definitively determine its full range and how common it is in comparison with the golden jackal, with which it can also interbreed. Frankfurt Zoological Society/www.guassaarea.org/www.fzs.org Below: Eric Didner Woollff! shouted Lajos Nemeth-Boka, lead naturalist and tour leader at GreenEye Ecotours. It was November of 2007. Nemeth-Boka was driving slowly along the west bank of the Nile River between Luxor and Aswan. An animal crossed the road in front of us, coming from the Nile s shore and running toward the Sahara sands, he says. I ve seen jackals and I ve seen wolves, and there is a big difference between the two. This was clearly a wolf. As it turned out, he was right. The first clues had come thousands of years earlier. But the wolf in jackal s clothing wasn t proved to be such until 2011, four years after Nemeth-Boka s sighting. In the fifth century BC, the Greek historian Herodotus reported that there were wolves in Egypt, describing them as no bigger than foxes. A century later, Aristotle reiterated that these Egyptian wolves were smaller than those of his homeland. And in the first century BC, the Greek historian Diodorus recounted a story that once a Nubian army, invading Egypt from the south, had been repelled by packs of wolves. The wolves drove the army all the way south from the town of Asyut to the thenborder, earning the town the Greek name of Lycopolis, city of the wolf.
In that ancient time, icons of the Egyptian wolf god Wepwawet were widespread. Wepwawet is a close relative of if not twin to the jackal god Anubis; both were the presiding deities of Lycopolis. Could Wepwawet, like Anubis, have had a living counterpart? Until recently, the only wolf universally recognized in Africa was the rare Ethiopian wolf (Canis simensis), found in the Ethiopian Highlands. The scientific consensus was that Egypt had no wolves, and that the ancient texts of Herodotus, Aristotle, and others must have referred to jackals. The Ethiopian wolf, a rare species, was until recently the only recognized wolf in Africa. It is not a subspecies of gray wolf. Rebecca Jackrel Even the Egyptian jackal, native to Egypt, Ethiopia, and Libya colloquially called the wolf-jackal was classified as a subspecies of the golden jackal: Canis aureus lupaster. But with a question mark, says ecologist Claudio Sillero-Zubiri, deputy director of the Wildlife Conservation Research Unit at the University of Oxford. What might the ancient Egyptians have known that we didn t? Millennia later, the true story is emerging. The tale picks up with an Indian biologist named Jugal K. Tiwari. A decade ago, Tiwari sent Sillero-Zubiri a picture from a video he had filmed in Eritrea. The footage showed a lanky canine with large paws that might have been a desert-
dwelling wolf, says Sillero-Zuburi. We hoped more information would turn up, but unfortunately it didn t. At least not right away. A golden jackal in Tanzania: Widespread in northern and central Africa, the species is also found in the Arabian Peninsula, eastern Europe, South and Central Asia, and Southeast Asia. Stefan Cook Then, while doing fieldwork in Ethiopia, biologists from universities in Addis Ababa, Ethiopia, and Oslo, Norway, noticed that certain golden jackals looked different from others. They were larger, more slender, and sometimes had a whitish color, says Nils Christian Stenseth, an ecologist and population biologist at the University of Oslo. The researchers collected scat specimens for mitochondrial DNA (mtdna) analysis. The samples, including some from more usual-looking golden jackals, were shipped to Stenseth s laboratory for analysis. The Oslo scientists soon alerted their counterparts on the project from other countries, including Sillero-Zubiri, that they had a rare find. The Egyptian jackal samples appeared to be wolf DNA. But when the team attempted to correlate them with other wolf samples in GenBank, the world s largest repository of genetic sequences, a surprise was in store. We could hardly believe our eyes when we found wolf DNA that didn t match anything in GenBank, says Eli K. Rueness, an evolutionary geneticist at the University of Oslo. Adds Sillero-Zubiri, We had unwittingly uncovered genetic evidence of a cryptic canid that looked like a golden jackal, but whose genetic code told another tale. In January 2011, Rueness, Stenseth, Sillero-Zubiri and colleagues unveiled their findings in the journal PLOS ONE: the Egyptian jackal is in fact a gray wolf. We now know that wolves were indeed in Africa in the days of the ancient Egyptians and long, long before, says Stenseth. Biologists have since updated the wolf s scientific name to Canis lupus lupaster, making it a subspecies of the gray wolf, C. lupus. The range of the gray wolf had been known to extend as far south as the Sinai Peninsula, in the subspecies C. lupus pallipes, the Indian wolf.
The Indian wolf, another subspecies of gray wolf, is found from India west into Saudi Arabia, but not in Africa. rahul sachdev But it didn t, it was thought, reach mainland Africa. Stenseth s team s results show that the lupaster subspecies inhabits at least two different locations in Ethiopia, some 1,500 miles southeast of Egypt. Phylogenetic reconstruction based on mtdna data suggests that Africa s gray wolf subspecies is most closely related to the Indian and Himalayan subspecies. Estimates using molecular clocks are that the ancestors of the African wolf split off from those of other gray wolves more than 2 million years ago and entered Africa by middle Pleistocene times, about 1 million years ago. That was before their northern cousins, the Eurasian wolves (C. lupus lupus), radiated across the northerly regions of the Earth, including the New World. After the 2011 revelation, biologist Philippe Gaubert of the Muséum National d Histoire Naturelle in Paris and colleagues spotted wolf-like animals on the periphery of packs of golden jackals near Kheune, in northwest Senegal, thousands of miles west of Egypt. The jackals turned out to be none other than lupaster. The wolves were larger and darker than golden jackals. They also behaved differently, with a solitary and somewhat shy demeanor. The only interactions between the two were when the wolves fought for carcasses eaten by golden jackals. The latter inevitably abandon[ed] their food to the former, noted Gaubert and his colleagues. The scientists looked at the mtdna of seven animals that proved to be lupaster: one from east of the Djoudj National Bird Sanctuary near Kheune; one from the Adrar des Ifoghas massif in eastern Mali; and five from Algeria s northeastern coastal region. The reassessed range of the African wolf from Ethiopia and Egypt across to Senegal, the researchers say, supports the idea of a wide spectrum of habitats for the species, from Mediterranean coastal and hilly areas (including hedged farmlands, scrublands, pinewoods and oak forests) in Algeria to tropical, semiarid savannas in Senegal and massifs in Mali. In 2012, the African wolf appeared again, triggering automatic cameras, or cameratraps, in Morocco s Middle Atlas Mountains, according to a group of zoologists led by Vicente Urios from University of Alicante in San Vicente del Raspeig, Spain. Without genetic analysis the identification is tentative, but if confirmed, it helps fill a gap in the
subspecies known distribution. Now biologists are asking: how many golden jackals across Africa are in fact African wolves? Lupaster looks like a large, blackish-yellow dog. Its tail is brushlike, with black hairs on the end. A mane of long, coarse, black-tipped fur runs from its crown to the base of its tail and onto its shoulders and hips. The golden jackal is smaller than lupaster, with soft, pale fur. Golden jackals are social animals: a breeding pair is often followed by its offspring, and the jackal sometimes forms packs when hunting. Its cry, heard just after dark or shortly before dawn, is a long, wailing howl, often followed by three yelps. In contrast, lupaster travels alone. A nocturnal creature, it s sometimes glimpsed as the Sun begins to set, when it emerges from caves and crevices, and from tombs. Whether it howls remains unknown. The field observations provide the means to distinguish the African wolf from the golden jackal. But hybridization between the two may be happening, at least in Senegal, based on detection of lupaster genes in C. aureus (golden jackals) there.
Cladogram, or branching family tree, of wolves and related species, is based on mitochondrial DNA (mtdna). Analysis of additional mtdna as well as nuclear DNA may modify details of this picture, but the African wolf, once considered a subspecies of golden jackal known as the Egyptian jackal, now falls firmly within the gray wolf group (Canis lupus). Steven R. Black/After E. K. Rueness et al., The Cryptic African Wolf, PLOS ONE 61 January 2011 Genetic techniques are revealing the hidden biodiversity of relatively unexplored places like Senegal and Ethiopia, lupaster s homelands. The understanding comes none too soon. Although golden jackals are listed by the International Union for Conservation of Nature (IUCN) as a species of least concern, lupaster may be much rarer. It s a priority, for both scientific and conservation efforts, to determine this wolf s whereabouts and numbers, says Stenseth. Gaubert s team agrees: Given that jackallike canids in Africa are regularly killed to protect livestock, it s urgent to [develop] a conservation strategy for the African wolf.
Lupaster s true identity shines a light on a formerly dark corner of the world, biogeographically, says Afework Bekele of Addis Ababa University. It s part of the Afroalpine fauna and flora, an assemblage of species that evolved in the relative isolation of the highlands of the Horn of Africa. Bekele is a member of the group that made the 2011 discovery of the wolf s genetic heritage. Ethiopia s Afroalpine highlands may hold the key to a better understanding of the new wolf. Lupaster has been seen most often in this land of short scrubby bushes sprouting from rock-strewn hillsides. Its haunts include the Menz Guassa Community Conservation Area (GCCA), some 160 miles northeast of Addis Ababa in the Menz- Gera Midir district. The GCCA is among the Ethiopian highlands most pristine and secluded natural wonders, says Zelealem Tefera, a scientist at the Frankfurt Zoological Society (FZS) Ethiopia Office. The FZS supports conservation projects throughout Africa in countries including Ethiopia, Tanzania, and Zambia. The Guassa area is managed through a common property resource system by people who live along its perimeter. The system traces to the seventeenth century, and is one of the oldest conservation management systems in sub-saharan Africa. Guassa communities live in farmers associations called kebeles. To generate an alternative income for these communities and to support management of the GCCA, ecotourism is being promoted by the FZS and by Ethiopian government agencies. Their efforts have led to the Guassa Community Lodge and several wilderness camping grounds. Visitors may experience a traditional village or a trek up steep hills to look for the Guassa plateau s rare species. The Ethiopian wolf and the gelada baboon are found only in the Ethiopian highlands. The Abyssinian hare, leopard, serval cat, civet, Egyptian mongoose, and spotted hyena, among others, share this mountaindesert. Lupaster walks among them. Out of the corner of your eye at sunset you might just spot an African wolf, says veterinarian Karen Laurenson, an ecologist and wildlife epidemiologist at the FZS- Ethiopia Office. She s glimpsed an animal that emerges at dusk, seemingly out of thin air, to return there just as quickly. I think I ve seen lupaster, but didn t know at the time what it was. Laurenson is concerned that the wolf could be extinct before we realize it. What we don t know, she says, is how many of these wolves there are. They may be widespread across Africa, but is that widespread in appreciable numbers, or dotted across certain habitats with only a few animals in each place? It s critical for their future that we find out. Small populations are prone to inbreeding, and may be quickly wiped out by diseases such as rabies and canine distemper virus. Disease and inbreeding aren t the only challenges lupaster faces. Uncovering this cryptic subspecies secrets may be a mixed blessing if, like the golden jackal, it lives near humans and snags farm animals. We know so little about this wolf, says Sillero-Zubiri. Who can say whether and when it takes sheep or other domesticated animals? It s still a shadow on a ridge. More than 80 percent of Guassa is highland, with eroded slopes that nonetheless support an expanding human population. Sheep are an important element of the farming system there and throughout northern Ethiopia, with their role increasing
because of the unreliability of crops. What do sheep graze on, in a brushy, highaltitude region such as Guassa? A prized natural resource Festuca grass (fescue). The inhabitants consider the grass their cloth, bread, and butter. One of the main reasons for protecting the Guassa area, FZS s Tefera says, is for harvesting good quality fescue. It s used for thatching, robes, and household implements. The grass is sold in distant markets in Addis Ababa and other cities. Guassa also provides a refuge for livestock when cultivated fields elsewhere lose their grasses, such as during droughts. Most of the sheep that regularly graze there come from adjacent villages. During prolonged droughts, however, sheep from villages farther away stay in Guassa in temporary pens to avoid long daily journeys. Therein lies the potential dust-up Lamenting that the golden ackal can be a pest and will attack domestic animals, one resident of Ethiopia asks, Why not this wolf? Similarly, shepherds in Senegal told Gaubert s team that the golden jackal was only observed preying on lambs, but that the African wolf may hunt larger livestock such as sheep, goats, and even cows. If lupaster indeed has a fondness for sheep and goats, adds Stenseth, that s bound to lead it into conflict with local agropastoral people. Luckily, says Tefera, the people here refer to it as the nomad jackal rather than the more common jackals they ve accused of killing their lambs. Shepherds in the Ethiopian Highlands know that jackals may take lambs; they do not know whether the newly identified wolf might be a threat to larger livestock. Ryan J. Burke/Armenz Guassa Community Conservation Area Its elusiveness may be lupaster s salvation. That, and a growing understanding by
people in places like Guassa that lupaster is a national heritage. It may also bring an income source through ecotourism but only for as long as it s with us, says Tefera. My Grandma told me about [these] wolves, offers an East African citizen, in response to the discovery of lupaster. I would ask if it was a hyena, dog, jackal, or fox, but her answer was, It s a wolf. She described it perfectly, too. She said back then they were common, but she hasn t seen any for decades. I ve heard many claims like that from older people. In ancient Egyptian art, the god Wepwawet was depicted as part human, part wolf, with the body of a human and the head of a wolf. Five thousands years later, will Wepwawet s incarnation as Africa s only gray wolf receive the recognition and protection it deserves? Scrapbook page from the travels, ca. 1891 1895, of William Vaughn Tupper includes a photograph of Asyut, on the Upper Nile, a city known to the ancient Greeks as Lycopolis, or City of the Wolf. Both the Egyptian wolf god, Wepwawet, and the jackal god, Anubis, were honored there. Their iconography is not always distinctive, but often a standing figure, such as the Anubis Tupper sketched, is likely to be Wepwawet. Part of the commentary reads: In the hills seen on the horizon are the tombs of the Priests and numberless holes in the rocks once filled with mummies of the Jackal.... The hills are now strewn with skulls and bones of the Jackals. Other nineteenth-century travelers referred to these as wolf mummies. William Vaughn Tupper Scrapbook Collection, Boston Public Library