Great Plains wolf

Last updated

Great Plains wolf
Face a face (28711329178).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
C. lupus
Subspecies:
C. l. nubilus
Trinomial name
Canis lupus nubilus
Say, 1823 [2]
Present distribution of wolf subspecies grtplains.jpg
Distribution of great plains wolf
Synonyms [3]

variabilis (Wied-Neuwied, 1841) [4]

The Great Plains wolf (Canis lupus nubilus), also known as the buffalo wolf or loafer, is a subspecies of gray wolf that once extended throughout the Great Plains, from southern Manitoba and Saskatchewan in Canada southward to northern Texas in the United States. [5] The subspecies was thought to be extinct in 1926, until studies declared that its descendants were found in Minnesota, Wisconsin and Michigan. [6] They were described as a large, light-colored wolf but with black and white varying between individual wolves, with some all white or all black. The Native Americans of North Dakota told of how only three Great Plains wolves could bring down any sized bison. [7]

Contents

Taxonomy

This wolf was first recorded in 1823 by the naturalist Thomas Say in his writings on Major Stephen Long's expedition to the Great Plains. Say was the first person to document the difference between a "prairie wolf" (coyote) and on the next page of his journal a wolf which he named Canis nubilus. He described one of these wolves that had been caught in a trap:

Canis nubilus. Dusky, the hair cinereous at base, then brownish-black then gray, then black; the proportion of black upon the hairs, is so considerable, as to give to the whole animal a much darker colour, than the darkest of the latrans, but the gray of the hairs combining with the black tips, in the general effect produce a mottled appearance; the gray colour predominates on the lower part of the sides; ears short, deep brownish-black, with a patch of gray hair on the anterior side within; muzzle blackish above; superior lifjs, anterior to the canine teeth, gray; inferior jaw at tip and extending in a narrowed line backwards, nearly to the origin of the neck, gray; beneath dusky ferruginous, greyish with long hair between the hind thighs, and with a large white spot on the breast; the ferruginous colour is very much narrowed on the neck, but is dilated on the lower part of the cheeks; legs brownish- black, with but a slight admixture of gray hairs, excepting on the anterior edge of the hind thighs, and the lower edgings of the toes, where the gray predominates; the tail is short, fusiform, a little tinged with ferruginous, black above near the base and at tip, the tip of the trunk hardly attaining to the os calcis; the longer hairs of the back, particularly over the shoulders, resemble a short sparse mane.....The aspect of this animal is far more fierce and formidable than either the common red wolf, or the prairie wolf, and is of a more robust form. [2]

In 1995, the American mammalogist Robert M. Nowak analyzed data on the skull morphology of wolf specimens from around the world. For North America, he proposed that there were only five subspecies of gray wolf. One of these he described as a moderate-sized wolf that was originally found from Texas to Hudson Bay and from Oregon to Newfoundland which he named C. l. nubilus. [8] [9] This proposal was not recognized in the taxonomic authority Mammal Species of the World (2005), which classified this wolf as one of the 27 subspecies of Canis lupus in North America. [3]

Lineage

"Roping gray wolf" - John C. H. Grabill photograph 1887 Grabill - Roping gray wolf.jpg
"Roping gray wolf" - John C. H. Grabill photograph 1887

Gray wolves (Canis lupus) migrated from Eurasia into North America 70,000–23,000 years ago [10] [11] and gave rise to at least two morphologically and genetically distinct groups. One group is represented by the extinct Beringian wolf [10] [12] and the other by the modern populations. [10]

A haplotype is a group of genes found in an organism that are inherited together from one of their parents. [13] [14] Mitochondrial DNA (mDNA) passes along the maternal line and can date back thousands of years. [15] A 2005 study compared the mitochondrial DNA sequences of modern wolves from across North America with those from thirty-four specimens dated between 1856 and 1915 collected from the western United States, Mexico and the Labrador Peninsula. The historic population was found to possess twice the genetic diversity of modern wolves, [16] [17] which suggests that the mDNA diversity of the wolves eradicated from the western US was more than twice that of the modern population. Some haplotypes possessed by the Mexican wolf, the extinct Southern Rocky Mountain wolf and the Great Plains wolf were found to form a unique "southern clade". All North American wolves group together with those from Eurasia, except for the southern clade which form a group exclusive to North America. The wide distribution area of the southern clade indicates that gene flow was extensive across the recognized limits of its subspecies. [17]

A study published in 2018 looked at the limb morphology of modern and fossil North American wolves. The major limb bones of the dire wolf, Beringian wolf, and most modern North American gray wolves can be clearly distinguished from one another. Late Pleistocene wolves on both sides of the Laurentide Ice Sheet — Cordilleran Ice Sheet possessed shorter legs when compared with most modern wolves. The Late Pleistocene wolves from the Natural Trap Cave, Wyoming and Rancho La Brea, southern California were similar in limb morphology to the Beringian wolves of Alaska. Modern wolves in the Midwestern USA and northwestern North America possess longer legs that evolved during the Holocene, possibly driven by the loss of slower prey. However, shorter legs survived well into the Holocene after the extinction of much of the Pleistocene megafauna, including the Beringian wolf. Holocene wolves from Middle Butte Cave (dated less than 7,600 YBP) and Moonshiner Cave (dated over 3,000 YBP) in Bingham County, Idaho were similar to the Beringian wolves. The Mexican wolf (C. l. baileyi) and pre-1900 samples of the Great Plains wolf (Canis lupus nubilus) resembled the Late Pleistocene and Holocene fossil gray wolves due to their shorter legs. [10]

Ancestor

In 2021, a mitochondrial DNA analysis of North American wolf-like canines indicates that the extinct Late Pleistocene Beringian wolf was the ancestor of the southern wolf clade, which includes the Mexican wolf and the Great Plains wolf. The Mexican wolf is the most ancestral of the gray wolves that live in North America today. [18]

Description

Buffalo Hunt, White Wolves Attacking Buffalo Bull - George Catlin 1844 George Catlin - Buffalo Hunt, White Wolves Attacking Buffalo Bull.jpg
Buffalo Hunt, White Wolves Attacking Buffalo Bull - George Catlin 1844

The Great Plains wolf's distribution once extended throughout the Great Plains from southern Manitoba and Saskatchewan southward to northern Texas. [5]

They are described as a large, light-colored wolf but with black and white varying between individual wolves, with some all white or all black. The average body length ranges from 1.4 m (4.6 ft) to 1.96 m (6.4 ft) [19] [20] with a weight of the male averaging 100 lb (45 kg) and the heaviest recorded at 150 lb (68 kg). The Native Americans of North Dakota told of how only three of these wolves could bring down a buffalo, including a large old bull. [7]

Early records indicate C. l. nubilus being very abundant throughout the Great Plains. After the mass extirpations of the American bison (Bison bison), they were poisoned and trapped for their pelts until few remained. The pioneer Alexander Henry wrote about these wolves several times during his trips to North Dakota, noting how they fed extensively on buffalo carcasses. They were bold around humans, sometimes approaching people and entering their tents while they slept. He recorded that Indians occasionally dug up wolf pups from their prairie dens and dug large pitfalls to capture wolves and foxes. Members of his group dug up wolf pups and found them very tame and easy to train. In 1833 Maximilian of Wied-Neuwied recorded that these wolves were common in the upper Missouri, where the Indians operated wolf pits and traded wolves to him in exchange for two rolls of tobacco each. He found the Indian's dogs to be more of a personal danger than the wolves. [7]

In 1856, Lt. G. K. Warren gathered together a collection of this wolf's skulls which now reside in the National Museum of Natural History. [7] He noted that some wolf skull specimens appeared not to be full-blooded wolves as their molars indicated a hybrid. There have been many stories in this region about ferocious hybrid wolf-dogs, and it is possible that the wolf's tameness and lack of fear of humans might be due admixture with domestic dogs. In North Dakota, by 1875 sightings of the wolf became rare, by 1887 they were almost gone. [7] On the Canadian Prairies, bounty payments for wolves commenced in 1878 in Manitoba, and 1899 in Saskatchewan and Alberta. [21] In North Dakota, two were sighted in 1915 by Remington Kellogg. The last known wolf was shot in 1922. [7] The Great Plains wolf was declared extinct in 1926. However, later studies found wolves in Minnesota, Wisconsin and Upper Michigan that were descendants of Canis lupus nubilus. Even then, their number became fewer and fewer until they were federally protected as an endangered species in 1974. Since then, their population became larger in the Great Lakes region and by 2009, their estimate grew to 2,992 wolves in Minnesota, 580 in Michigan and 626 in Wisconsin. A R [22] [19] [20] [6]

See also

Related Research Articles

Subspecies of <i>Canis lupus</i>

There are 38 subspecies of Canis lupus listed in the taxonomic authority Mammal Species of the World. These subspecies were named over the past 250 years, and since their naming, a number of them have gone extinct. The nominate subspecies is the Eurasian wolf.

<span class="mw-page-title-main">Red wolf</span> Canid native to the southeastern United States

The red wolf is a canine native to the southeastern United States. Its size is intermediate between the coyote and gray wolf.

<span class="mw-page-title-main">Dire wolf</span> Extinct species of the genus Aenocyon from North America

The dire wolf is an extinct canine. The dire wolf lived in the Americas during the Late Pleistocene and Early Holocene epochs. The species was named in 1858, four years after the first specimen had been found. Two subspecies are recognized: Aenocyon dirus guildayi and Aenocyon dirus dirus. The largest collection of its fossils has been obtained from the Rancho La Brea Tar Pits in Los Angeles.

<i>Canis</i> Genus of carnivores

Canis is a genus of the Caninae which includes multiple extant species, such as wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and dentition, long legs, and comparatively short ears and tails.

<span class="mw-page-title-main">Eastern wolf</span> Subspecies of carnivore

The eastern wolf also known as the timber wolf, Algonquin wolf or eastern timber wolf, is a canine of debated taxonomy native to the Great Lakes region and southeastern Canada. It is considered to be either a unique subspecies of gray wolf or red wolf or a separate species from both. Many studies have found the eastern wolf to be the product of ancient and recent genetic admixture between the gray wolf and the coyote, while other studies have found some or all populations of the eastern wolf, as well as coyotes, originally separated from a common ancestor with the wolf over 1 million years ago and that these populations of the eastern wolf may be the same species as or a closely related species to the red wolf of the Southeastern United States. Regardless of its status, it is regarded as unique and therefore worthy of conservation with Canada citing the population in eastern Canada as being the eastern wolf population subject to protection.

<span class="mw-page-title-main">Hokkaido wolf</span> Extinct subspecies of the gray wolf

The Hokkaido wolf, also known as the Ezo wolf and in Russia as the Sakhalin wolf, is an extinct subspecies of gray wolf that once inhabited coastal northeast Asia. Its nearest relatives were the wolves of North America rather than Asia. It was exterminated in Hokkaido during the Meiji Restoration period, when American-style agricultural reforms incorporated the use of strychnine-laced baits to kill livestock predators. Some taxonomists believe that it survived up until 1945 on the island of Sakhalin. It was one of two subspecies that were once found in the Japanese archipelago, the other being the Japanese wolf.

<span class="mw-page-title-main">Mexican wolf</span> Subspecies of grey wolf

The Mexican wolf, also known as the lobo, is a subspecies of gray wolf native to southeastern Arizona and southern New Mexico in the United States, and northern Mexico. It once also ranged into western Texas. It is the smallest of North America's gray wolves, and is similar to the Great Plains wolf, though it is distinguished by its smaller, narrower skull and its darker pelt, which is yellowish-gray and heavily clouded with black over the back and tail. Its ancestors were likely the first gray wolves to enter North America after the extinction of the Beringian wolf, as indicated by its southern range and basal physical and genetic characteristics.

<span class="mw-page-title-main">Italian wolf</span> Subspecies of carnivore

The Italian wolf, also known as the Apennine wolf, is a subspecies of the grey wolf native to the Italian Peninsula. It inhabits the Apennine Mountains and the Western Alps, though it is undergoing expansion towards the north and east. As of 2022 the wolf population within Italy is estimated to be 3,307 individuals. Although not universally recognised as a distinct subspecies, it nonetheless possesses a unique mtDNA haplotype and a distinct skull morphology.

<span class="mw-page-title-main">Northwestern wolf</span> Subspecies of mammal

The northwestern wolf, also known as the Mackenzie Valley wolf, Alaskan timber wolf, or Canadian timber wolf, is a subspecies of gray wolf in western North America. Arguably the largest gray wolf subspecies in the world, it ranges from Alaska, the upper Mackenzie River Valley; southward throughout the western Canadian provinces, aside from prairie landscapes in its southern portions, as well as the Northwestern United States.

<span class="mw-page-title-main">Indian wolf</span> Wolf subspecies

The Indian wolf is a subspecies of gray wolf that ranges from Southwest Asia to the Indian subcontinent. It is intermediate in size between the Himalayan wolf and the Arabian wolf, and lacks the former's luxuriant winter coat due to it living in warmer conditions. Within this subspecies, the "Indian plains wolf" is genetically basal to all other extant Canis lupus apart from the older-lineage Himalayan wolf, with both proposed as separate species. The Indian wolf travels in smaller packs and is less vocal than other variants of the gray wolf, and has a reputation for being cunning. The Indian wolf is one of the most endangered populations of gray wolf in the world.

<span class="mw-page-title-main">Japanese wolf</span> Extinct subspecies of the gray wolf

The Japanese wolf, also known as the Honshū wolf, is an extinct subspecies of the gray wolf that was once endemic to the islands of Honshū, Shikoku and Kyūshū in the Japanese archipelago.

<span class="mw-page-title-main">Himalayan wolf</span> Subspecies of mammal

The Himalayan wolf is a canine of debated taxonomy. It is distinguished by its genetic markers, with mitochondrial DNA indicating that it is genetically basal to the Holarctic grey wolf, genetically the same wolf as the Tibetan and Mongolian wolf, and has an association with the African wolf. No striking morphological differences are seen between the wolves from the Himalayas and those from Tibet. The Himalayan wolf lineage can be found living in Ladakh in the Himalayas, the Tibetan Plateau, and the mountains of Central Asia predominantly above 4,000 m (13,000 ft) in elevation because it has adapted to a low-oxygen environment, compared with other wolves that are found only at lower elevations.

<span class="mw-page-title-main">Southern Rocky Mountain wolf</span> Extinct subspecies of the gray wolf

The southern Rocky Mountain wolf is an extinct subspecies of gray wolf which was once distributed over southeastern Idaho, southwestern Wyoming, northeastern Nevada, Utah, western and central Colorado, northwestern Arizona, and northwestern New Mexico. It was a light-colored, medium-sized subspecies closely resembling the Great Plains wolf, though larger, with more blackish-buff hairs on the back. This wolf went extinct by 1935. Wolves of the subspecies Canis lupus occidentalis have now been reestablished in Idaho and Wyoming.

<span class="mw-page-title-main">Beringian wolf</span> Extinct type of wolf that lived during the Ice Age in Alaska, Yukon, and northern British Columbia

The Beringian wolf is an extinct population of wolf that lived during the Ice Age. It inhabited what is now modern-day Alaska, Yukon, and northern British Columbia. Some of these wolves survived well into the Holocene. The Beringian wolf is an ecomorph of the gray wolf and has been comprehensively studied using a range of scientific techniques, yielding new information on their prey species and feeding behaviors. It has been determined that these wolves are morphologically distinct from modern North American wolves and genetically basal to most modern and extinct wolves. The Beringian wolf has not been assigned a subspecies classification and its relationship with the extinct European cave wolf is not clear.

<span class="mw-page-title-main">Pleistocene coyote</span> Extinct subspecies of carnivore

The Pleistocene coyote, also known as the Ice Age coyote, is an extinct subspecies of coyote that lived in western North America during the Late Pleistocene era. Most remains of the subspecies were found in southern California, though at least one was discovered in Idaho. It was part of a North American carnivore guild that included other canids like foxes, gray wolves, and dire wolves. Some studies suggest that the Pleistocene "coyote" was not in fact a coyote, but rather an extinct western population of the red wolf.

<span class="mw-page-title-main">Pleistocene wolf</span> Extinct lineage of the grey wolf

The Pleistocene wolf, also referred to as the Late Pleistocene wolf, is an extinct lineage or ecomorph of the grey wolf. It was a Late Pleistocene 129 Ka – early Holocene 11 Ka hypercarnivore. While comparable in size to a big modern grey wolf, it possessed a shorter, broader palate with large carnassial teeth relative to its overall skull size, allowing it to prey and scavenge on Pleistocene megafauna. Such an adaptation is an example of phenotypic plasticity. It was once distributed across the northern Holarctic. Phylogenetic evidence indicates that despite being much smaller than the prehistoric wolf, the Japanese wolf, which went extinct in the early 20th century, was of a Pleistocene wolf lineage, thus extending its survival to several millennia after its previous estimated extinction around 7,500 years ago.

<span class="mw-page-title-main">Cave wolf</span> Extinct Ice-Age European subspecies of wolf

The cave wolf is an extinct glacial mammoth steppe-adapted white wolf that lived during the Middle Pleistocene to the Late Pleistocene. It inhabited Europe, where its remains have been found in many caves. Its habitat included the mammoth steppe grasslands and boreal needle forests. This large wolf was short-legged compared to its body size, yet its leg size is comparable with that of the Arctic wolf C. l. arctos. Mitochondrial DNA analysis shows it to be more closely related to the domestic dog than the modern wolf, indicating possible ancestry.

<span class="mw-page-title-main">Evolution of the wolf</span>

The evolution of the wolf occurred over a geologic time scale of at least 300 thousand years. The grey wolf Canis lupus is a highly adaptable species that is able to exist in a range of environments and which possesses a wide distribution across the Holarctic. Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other. This variation in sub-populations is closely linked to differences in habitat – precipitation, temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.

<i>Canis mosbachensis</i> Extinct species of carnivore

Canis mosbachensis is an extinct wolf that once inhabited Europe 600,000—420,000 years ago. The Mosbach wolf was a short-legged carcass feeder adapted for scavenging megafauna on the mammoth steppe. The Mosbach wolf is proposed as the ancestor of the grey wolf Canis lupus but some mammalogists have assigned it as the subspecies Canis lupus mosbachensis.

References

  1. ↑ "Canis lupus nubilus". Committee on the Status of Endangered Wildlife in Canada (COSEWIC): Not at Risk (4/1/1999)
  2. 1 2 Say, T. et al. (1823) Account of an expedition from Pittsburgh to the Rocky Mountains, performed in the years 1819 and '20 : by order of the Hon. J.C. Calhoun, sec'y of war: under the command of Major Stephen H. Long. From the notes of Major Long, Mr. T. Say, and other gentlemen of the exploring party . Philadelphia : H.C. Carey and I. Lea ... p. 169-173.
  3. 1 2 Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 575–577. ISBN   978-0-8018-8221-0. OCLC   62265494. url=https://books.google.com/books?id=JgAMbNSt8ikC&pg=PA576
  4. ↑ Prince Maximilian of Wied-Neuwied (1841). Reise in das innere Nord-Amerika in den Jahren 1832 bis 1834. Vol. 2. Coblenz. p. 95.[Trip to the interior of North America 1832 to 1834]
  5. 1 2 Mech, L. (1970). "Appendix A – Subspecies of wolves – North American". The Wolf: The Ecology and Behavior of an Endangered Species. Doubleday. ISBN   978-0-307-81913-0. Great Plains wolf; buffalo wolf; loafer. This is another extinct subspecies. It once extended throughout the Great Plains from southern Manitoba and Saskatchewan southward to northern Texas.
  6. 1 2 Wydeven, Adrian P; Van Deelen, Timothy R; Heske, Edward J, eds. (2009). Recovery of Gray wolves in the Great Lakes Region of the United States, An Endangered Subpecies Success Story. link.springer.com. doi:10.1007/978-0-387-85952-1. ISBN   978-0-387-85951-4. S2CID   132793403.
  7. 1 2 3 4 5 6 A Biological Survey of North Dakota, Vernon, B., (1926), North American Fauna: Number 49: pp. 150–156.
  8. ↑ Nowak, R. (2003). "9-Wolf Evolution and Taxonomy". In Mech, L. David; Boitani, Luigi (eds.). Wolves: Behaviour, Ecology and Conservation. University of Chicago Press. p. 245. ISBN   0-226-51696-2.
  9. ↑ Another look at wolf taxonomy Nowak, R.M. 1995. Pp. 375–397 in L.N. Carbyn, S.H. Fritts and D.R. Seip, eds. Ecology and conservation of wolves in a changing world: proceedings of the second North American symposium on wolves. Canadian Circumpolar Institute, University of Alberta, Edmonton, Canada.
  10. 1 2 3 4 Tomiya, Susumu; Meachen, Julie A (2018). "Postcranial diversity and recent ecomorphic impoverishment of North American gray wolves". Biology Letters. 14 (1): 20170613. doi:10.1098/rsbl.2017.0613. PMC   5803591 . PMID   29343558.
  11. ↑ Koblmüller, Stephan; Vilà, Carles; Lorente-Galdos, Belen; Dabad, Marc; Ramirez, Oscar; Marques-Bonet, Tomas; Wayne, Robert K.; Leonard, Jennifer A. (2016). "Whole mitochondrial genomes illuminate ancient intercontinental dispersals of grey wolves (Canis lupus)". Journal of Biogeography. 43 (9): 1728–1738. doi:10.1111/jbi.12765. S2CID   88740690.
  12. ↑ Leonard, J. A.; Vilà, C; Fox-Dobbs, K; Koch, P. L.; Wayne, R. K.; Van Valkenburgh, B (2007). "Megafaunal extinctions and the disappearance of a specialized wolf ecomorph" (PDF). Current Biology. 17 (13): 1146–50. doi:10.1016/j.cub.2007.05.072. hdl: 10261/61282 . PMID   17583509. S2CID   14039133. Archived from the original (PDF) on 2016-12-28. Retrieved 2018-01-27.
  13. ↑ Cox, C. B.; Moore, Peter D.; Ladle, Richard (2016). Biogeography: An Ecological and Evolutionary Approach. Wiley-Blackwell. p. 106. ISBN   978-1-118-96858-1.
  14. ↑ Editorial Board (April 2012). Concise Dictionary of Science. V&s Publishers. ISBN   978-93-81588-64-2.
  15. ↑ Arora, Devender; Singh, Ajeet; Sharma, Vikrant; Bhaduria, Harvendra Singh; Patel, Ram Bahadur (2015). "Hgs Db: Haplogroups Database to understand migration and molecular risk assessment". Bioinformation. 11 (6): 272–5. doi:10.6026/97320630011272. PMC   4512000 . PMID   26229286.
  16. ↑ Miklosi, Adam (2015). Dog Behaviour, Evolution, and Cognition. Oxford Biology (2 ed.). Oxford University Press. pp. 106–107. ISBN   978-0199545667.
  17. 1 2 Leonard, Jennifer A.; Vilà, Carles; Wayne, Robert K. (2004). "FAST TRACK: Legacy lost: Genetic variability and population size of extirpated US grey wolves (Canis lupus)". Molecular Ecology. 14 (1): 9–17. doi:10.1111/j.1365-294X.2004.02389.x. PMID   15643947. S2CID   11343074.
  18. ↑ Wilson, Paul J.; Rutledge, Linda Y. (2021). "Considering Pleistocene North American wolves and coyotes in the eastern Canis origin story". Ecology and Evolution. 11 (13): 9137–9147. doi:10.1002/ece3.7757. PMC   8258226 . PMID   34257949.
  19. 1 2 "Great Plains Wolf". www.wolfquest.org.
  20. 1 2 "Great Plains Wolf". wolf-stuff.com.
  21. ↑ Proulx, Gilbert; Rodtka, Dwight (2015). "Predator Bounties in Western Canada Cause Animal Suffering and Compromise Wildlife Conservation Efforts". Animals. 5 (4): 1034–1046. doi: 10.3390/ani5040397 . PMC   4693201 . PMID   26479482.
  22. ↑ "The Great Plains Wolf". www.cosmosmith.com.

Wikispecies-logo.svg Data related to Canis lupus nubilus at Wikispecies

This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.