The Tibetan wolf was found to be the most highly divergent of the Old World wolves, had suffered a historical population bottleneck and had only recently recolonized the Tibetan Plateau. Tibet).  Therefore, haplogroup 1 was able to enter into North America during this period. The phylogentic tree generated from the sequences showed that four of the Siberian canids dated 28,000 YBP and one Canis c.f. Rather, all living wolves in North America today originated from one colonisation event, researchers report in the Journal of Biogeography. Ph.D. dissertation", "Rethinking dog domestication by integrating genetics, archeology, and biogeography", https://en.wikipedia.org/w/index.php?title=Evolution_of_the_wolf&oldid=992501429, Articles lacking reliable references from September 2016, Wikipedia articles needing page number citations from July 2016, Creative Commons Attribution-ShareAlike License, Proposed evolution and branching of genus. This has caused a dramatic decrease in its population size over the last two centuries. , Comparison to the gray wolf lineage indicated that Taimyr-1 was basal to gray wolves from the Middle East, China, Europe and North America but shared a substantial amount of history with the present-day gray wolves after their divergence from the coyote. variabilis group that were a comparable size to the extant Indian wolf (Canis lupus pallipes), and the large hypercarnivorous Canis (Xenocyon) lycaonoides that was comparable in size to extant northern grey wolves.  The replacement in North America of a basal population of wolves by a more recent one supported the findings of earlier studies. Idaho wolf reintroduction was less of a national issue- The reintroduction of the wolf to central Idaho was overshadowed in the mind of the public by the greater prominence of the Yellowstone wolf restoration project. In North America it gave rise to early Canis which first appeared in the Miocene (6 million YBP) in south-western US and Mexico.  This variation in sub-populations is closely linked to differences in habitat – precipitation, temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.  "Dogs are not paedomorphic wolves. Wolves also have serological and biochemical traits distinct from dogs (Leone and Wiens 1956; Lauer, Kuyt & Baker 1969). This new finding could change conservation plans. Past studies based on SNPs, genome-wide similarities with Chinese wolves, and lower linkage disequilibrium might reflect regional admixture between dogs with wolves and gene flow between dog populations, with divergent dog breeds possibly maintaining more wolf ancestry in their genome. The study proposed that analysis of ancient DNA might be a better approach. in Latin means confer, uncertain) from Italy dated to 2.2 million YBP. lepophagus. lupus. The wolves, however, might come in sheep's clothing (Matthew 7:15), and so the false prophets, the usurpers of authority, and leaders of parties within the Church, are also included in the term. However, much of their diversity was later lost during the twentieth century. , In 2013, a genetic study found that the wolf population in Europe was divided along a north–south axis and formed five major clusters. & Wessels, W. The carbon dating gave a calendar-year age estimate that ranged between 16,945–13,905 YBP. The time between generations for wolves is assumed to be three years based on the extant gray wolf, and two years for the dog based on the extant dog. They exhibited a lower percentage of tooth breakage comparable with MIS-3 wolves. Coyotes derived from an exclusively North America lineage of the genus. The wolves, too, were closely related to each other than to the dogs. , In 2016, a study investigated for the first time the population subdivisions, demography, and the relationships of gray wolves based on their whole-genome sequences. The study excluded the sequences of the divergent Himalayan wolf and the Indian gray wolf. Another study based on SNPS showed wolves clustering with coyotes and separate from dogs. There is evidence of gene flow between African golden wolves, golden jackals, and gray wolves. They were encouraged to recalculate their divergence dates accordingly. The fossil evidence suggests that grey wolves were abundant in North America as long as 500,000 years ago. ): Contributions in Quaternary Vertebrate Paleontology: A Volume in Memorial to John E. Guilday, 218–227.  This was followed by an explosion of Canis evolution across Eurasia in the Early Pleistocene around 1.8 million YBP in what is commonly referred to as the wolf event. The oldest Canis remains found in Europe were from France and dated to 3.1 million YBP, followed by Canis cf. By the Late Holocene, the dog's niche area was less in size than researchers had expected to find, indicating that it was limited by biotic factors. :p241 However, there are no fossils of C. rufus until the Late Rancholabrean. Closely related to this haplotype was one that was found in the recently extinct Japanese wolf. This was surprising. They developed strong jaws and the highest flesh-slicing ability compared to the other wolves, with shallower jaws than the modern wolf but broader and deeper jaws than MIS 7 and MIS 5 wolves. Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other. A VOLUME IN HONOUR OF HANS DE BRUIJN – DEINSEA 10: 563–574 [ISSN 0923-9308] Published 1 December 2003. Bears, deer and people followed, but the exact timing of the influx is unclear. Lawrence, B. , In 2012, a study of the wolf-like Canis species of ancient China under the direction of Xiaoming Wang found that these were all quite close to C. lupus in both dental and post-cranial dimensions except for Canis variabilis, which was "very strange" compared to other Canis in China as it had much smaller cranio-dental dimensions than earlier and later species. :181, Merriam named 3 unusual species based on specimens recovered from the Rancho La Brea tar pits. Canis dirus lived in the late Pleistocene to early Holocene in North and South America  and was the largest of all Canis species. :p240, Canis lepophagus lived in the early Pliocene in North America. Paleoenvironment was cold, open tundra with summer temperatures between 7 °C and 11 °C and winter temperatures between −10 °C and −30 °C dominated by reindeer and bison. Since wolf recovery plans were first written in the 1980s, people have learned much more about wolves’ behavior, ecology and needs. All other haplotypes were basal in the tree, and these formed 2–3 smaller clades that were assigned to haplogroup 2 that was not monophyletic.. The taxonomic status of uncertain species is better resolved through using nuclear DNA from the nucleus of the cell, which is more suitable for analyzing the recent history. This implies that the original wolf populations were out-competed by a new type of wolf which replaced them. The studies found that precipitation and mean diurnal temperature range were the most influential variables on sequence variation. The Italian wolf underwent a reduction in body size with the loss of the red deer in Italy during the Renaissance.  A 2004 study that used 310 wolf skulls and over 700 dog skulls representing 100 breeds concluded that the evolution of dog skulls can generally not be described by heterochronic processes such as neoteny although some pedomorphic dog breeds have skulls that resemble the skulls of juvenile wolves. This introgression could have provided early dogs living in high latitudes with phenotypic variation beneficial for adaption to a new and challenging environment. :p60 The large wolf C. antonii from late Pliocene to early Pleistocene China was assessed as being a variation within C. chihliensis,:p197 and the large wolf C. falconeri occurred abruptly in Europe in the Early Pleistocene, perhaps representing a westward extension of C. Moreover, domestic dogs occupy a range of novel shapes outside the domain of wild carnivorans.  The tree illustrates the genotype-phenotype distinction, where a genotype is an organism's full hereditary information and a phenotype is an organism's actual observed properties, such as morphology, development, or behavior. , The dingo (Canis lupus dingo) refers to the dog found in Australia. For the ancient wolves of North America, instead of an extinction/replacement model suggested by a previous study, this study found substantial evidence of a population bottleneck in North America in which the ancient wolf diversity was almost lost around the beginning of the Holocene (no further elaboration in the study). , In 2016, two studies compared the sequences of 42,000 single nucleotide polymorphisms in North American gray wolves and found that they formed six ecotypes. , There are two key assumptions that are made for dating the divergence time for species: the generation time and the genetic mutation rate per generation.  The first definite wolf appeared in the Late Blancan/Early Irvingtonian,:p240 and named C. priscolatrans that was either very close to or a synonym for Canis edwardii. , In 2015, a study looked at the paleoecology of large carnivores across the Mammoth steppe during the Late Pleistocene by using stable isotope analysis of their fossil collagen to reconstruct their diets. "The difference in size and proportion between some breeds are as great as those between any wild genera, but all dogs are clearly members of the same species. :p148:p181 C. chihliensis appears to be more primitive and smaller than C. lupus, and measurements of its skull and teeth are similar to C. lupus but those of its postcranial elements are smaller. The study also indicated that all North America wolves have a significant amount of coyote ancestry and all coyotes some degree of wolf ancestry, and that the red wolf and eastern wolf are highly admixed with different proportions of gray wolf and coyote ancestry. The gray wolf, jackals, and hunting dogs derived from the Eurasian lineage. "We showed all of the genetic diversity, including the Mexican wolves, is compatible with a single colonisation event from the Eurasian population," says Leonard. Filming the ''Dances With Wolves'' stampede -- Producer Jim Wilson shares how the dangerous scene was created with the help of Neil Young, Oreos, and a lot of real buffalo  Kurten proposed that the Blancan C. lepophagus derived from smaller Miocene Canis species in North America. , The remains of the larger coyote-like Canis edwardii have been found in the later Pliocene in the south-western USA along with C. lepophagus, which indicates a descent. , Canis occidentalis furlongi (Merriam 1910) was described as a wolf considerably smaller than the dire wolf and more closely related to the timber wolf Canis lupus occidentalis. These six wolf ecotypes were named West Forest, Boreal Forest, Arctic, High Arctic, Baffin, and British Columbia.  Nowak proposed the name Canis lupus furlongi as he believed that it was a subspecies of the gray wolf. Hall and H.S. In Eurasia, the loss of ancient lineages could not be simply explained and appears to have been slow across time with the reasons unclear. WOLVES ARE CRUCIAL FOR BIODIVERSITY. lupus. The mammals of the Huachuca Mountains, southeastern Arizona. The wolves in the study formed 3 clades: north Asian wolves that included those from northern China and eastern Russia, Himalayan wolves from the Tibetan Plateau, and a unique population from southern China. Phylogenetic relationship between four canids. The separation between dog and wolf may reflect the rapid rate in which domestication occurred, including the possibility of a second domestication event occurring in North America. In Asia, only four haplotypes have been identified as belonging to this haplogroup, and two of them occur in the Middle East. In 2018, whole genome sequencing was used to compare members of genus Canis, along with the dhole (Cuon alpinus) and the African hunting dog (Lycaon pictus). This analysis suggests that the Pleistocene wolves from haplogroup 2 found in Beringia and Belgium preyed mainly on Pleistocene megafauna, which became rare at the beginning of the Holocene 12,000 years ago. These results indicate that a historic population of several hundred thousand wolves once existed in Mexico and the western US. Muséum national d'Histoire naturelle, Paris, 334 p. (Mémoires du Muséum national d'Histoire naturelle, Sér. The study indicated that there exists individual wolves of dog/wolf ancestry in most of the wolf populations of Eurasia but less so in North America. They improve habitat and increase populations of countless species from birds of prey to pronghorn, and even trout.  These characteristics of shortened rostrum, tooth crowding, and absence or rotation of premolars have been documented in both ancient and modern wolves. These responses to dietary changes showed species-wide dietary shifts, and not just local ecomorphs, in response to climatic and ecological variables. Surprisingly, the Mexican wolf was long believed to have its own origin. Researchers are limited to morphologic analysis but it is difficult to estimate the intra-species and inter-species variations and relationships that existed between specimens across time and place. After the Ice Age, the grey wolves were well adapted to the environment and prey that was present, so they were free to dominate. , The reduced body size of the early dog compared to a wolf is thought due to niche selection (Olsen 1985; Morey 1992; Coppinger & The gray wolf is thought to be ancestral to the domestic dog, however its relationship to C. variabilis, and the genetic contribution of C. variabilis to the dog, is the subject of debate. They exhibited a higher percentage of tooth breakage and severely worn teeth compared to the other wolves, and may have been using their upper P4 and lower m1 to crush bone rather than their molars, leading to a higher frequency of damage. "Now they've become inbred, so they have lots of genetic problems," says Leonard. , In 2016, a study built on the work of another major study and analyzed the sequences of 12 genes that are located on the heavy strand of the mitochondrial genome of extinct and modern C. lupus. A scenario consistent with the phylogenetic, ice sheet and sea-level data was that during the Ice Age when sea levels were at their lowest, there was a single wave of wolf colonization into North America starting with the opening of the Bering land bridge 70,000 YBP and closing during the Late Glacial Maximum of the Yukon corridor that ran through the division between the Laurentide Ice Sheet and the Cordilleran Ice Sheet 23,000 YBP. Lower sea levels meant that a land bridge provided a direct route connecting the two continents, before the end of the last Ice Age some 12,000 years ago. The techniques used to extract, locate and compare genetic sequences can be applied using advances in technology, which allows researchers to observe longer lengths of base pairs that provide more data to give better phylogenetic resolution.  In 2016, a whole-genome DNA study proposed, based on the assumptions made, that all of the North American wolves and coyotes diverged from a common ancestor less than 6,000–117,000 years ago. , There is little genetic information available on the ancient wolves that existed prior to the bottleneck. It’s believed that the word “werewolf” came around the 5th century, derived from the Old English “werewulf”.  Paleolithic dogs were directly associated with human hunting camps in Europe over 30,000 (YBP) and it is proposed that they were domesticated.  The phylogenetic descent of the extant wolf C. lupus from C. etruscus through C. mosbachensis is widely accepted. When Did Wild Wolves Become Pet Dogs?  The past sea levels of the Korean Strait together with the timing of the Japanese wolf sequences indicated that it arrived to the southern islands less than 20,000 YBP. From the tree, the study was able to infer the most recent common ancestor for all other C. lupus specimens – modern and extinct – was 80,000 YBP and this date concurred with the earlier study. While it is unlikely that grey wolves contributed to the dire wolf's extinction, there are several possible reasons why the less ferocious of the two survived.  The Pleistocene Eurasian wolves have been found to be morphologically and genetically comparable to the Pleistocene eastern-Beringian wolves, with some of the ancient European and Beringian wolves sharing a common haplotype (a17), which makes ecological similarity likely.  Over the past million years, numerous wolf-like forms existed but their turnover has been high, and modern wolves are not the lineal ancestors of dogs. Basilosaurus did share some traits with marine reptiles, ... Often called “wolves with hooves,” mesonychids were medium- to large-sized predators … They show a phylogenetic relationship with extirpated wolves from the south (Oklahoma), indicating that these wolves are the last remains of a once widespread group that has been largely extirpated during the last century, and that the wolves of northern North America had originally expanded from southern refuges below the Wisconsin glaciation after the ice had melted at the end of the Last Glacial Maximum. Do dogs descend from wolves?  At the end of the most recent glacial retreat during the past 30,000 years, warming melted the glacial barriers across northern Canada allowing arctic mammals to extend their range into mid-latitude North America, including elk, caribou, bison, and the gray wolf. This was followed by a single population of modern wolves expanding out of a Beringia refuge to repopulate the wolf's former range, replacing the remaining Late Pleistocene wolf populations across Eurasia and North America as they did so.  The archaeological and paleontological records show their continuous presence for at least the last 300,000 years. The study suggests that the African golden wolf is a descendant of a genetically admixed canid of 72% grey wolf and 28% Ethiopian wolf ancestry, and that the Ethiopian wolf once had a wider range in Africa.  Wolves went through a population bottleneck 20,000 YBP that coincides with the Last Glacial Maximum, which indicates that many wolf populations may have gone extinct at the same time as the Beringian wolves. It was therefore assumed that today's wolves are descended from these ancient predators.  The study suggested that previous datings based on the divergence between wolves and coyotes of one million years ago using fossils of what appeared to be coyote-like specimens may not reflect the ancestry of the modern forms. , The modern gray wolf expanded out of Beringia 25,000 years ago. When Yellowstone National Park was created in 1872, gray wolf (Canis lupus) populations were already in decline in Montana, Wyoming and Idaho. , The finding of a second wolf specimen from the same area (“Taimry-2”) and dated to 42,000 YBP has also been sequenced but yielded only mitochondrial DNA. Its skull and dentition were described as being intermediate between Canis lupus occidentalis and the dire wolf. Elk move into heavy timber when wolves are around, Creel added, but return to the grassy, open meadows when wolves go away. :p242 during the Middle Pleistocene in North America. , The dog was a very successful invader of North America and had established a widespread ecological niche by the Early–Middle Holocene. However, its premolar P4 (upper carnassials) were massive, and the hypocone of the molar M1 was larger than that of the dire wolf.  Such an early divergence is consistent with several paleontological reports of dog-like canids dated up to 36,000 YBP, as well as evidence that domesticated dogs most likely accompanied early colonizers into the Americas.  Recent mitochondrial DNA analyses of ancient and modern gray wolf specimens supports a pattern of population reduction and turnover. He noted that C. arnensis of Europe showed striking similarities to C. priscolatrans, and they could represent what once was a holarctic population of coyotes.