caninae

{{Cladogram

|align=left|title=Canid subfamilies

|cladogram={{clade|style=font-size:85%;line-height:80%;width:262px;

|style1=background-color:#ccccff;

|label1=Canidae

|1={{clade

|1={{extinct}}Hesperocyoninae 80 px

|2={{clade

|1={{extinct}}Borophaginae 70 px

|2= Caninae 70 px

}}

}}

}}

}}

The genus Leptocyon (Greek: leptos slender + cyon dog) includes 11 species and was the first primitive canine. They were small and weighed around 2 kg.{{rp|page=53}} They first appeared in Sioux County, Nebraska in the Orellan era 34-32 million years ago, which was the beginning of the Oligocene. This was the same time as the appearance of the Borophaginae with whom they share features, indicating that these were two sister groups. Borophaginae skull and dentition were designed for a powerful killing bite compared with the Leptocyon which were designed for snatching small, fast-moving prey. The species L. delicatus is the smallest canid to have existed. At the close of their genus 9 million years ago one Leptocyon lineage resembled the modern fox.{{rp|page=53}} The various species of Leptocyon branched 11.9 Mya into Vulpini (foxes) and Canini (canines).{{rp|pages=174–175}}

The canines spent two-thirds of their history in North America, before dispersing 7 million years ago into Asia, Europe, and Africa. One of the characteristics that distinguished them from the Borophaginae and Hesperocyoninae was their possession of less weight in their limbs and more length in their legs, which may have aided their dispersion. The first canine to arrive in Eurasia was the coyote-sized Canis cipio, whose scant fossils were found in Spain. However, the assignment of C. cipio within the canines to the genus Canis or genus Eucyon is not clear.{{rp|pages=143–144}}

The results of allozyme and chromosome analyses have previously suggested several phylogenetic divisions:

class="wikitable collapsible"
Divisions ! Description ! Image ! Genus ! Species
rowspan="4" style="text-align:center;"| Subtribe Canina ! rowspan="4" style="text-align:center;"|The wolf and wolf-like canines | 150px |Canis {{small|Linnaeus, 1758}} | Canis aureus Canis familiaris Canis latrans Canis lupaster Canis lupus Canis rufus Canis lycaon Canis simensis † Canis antonii † Canis armbrusteri † Canis chihliensis † Canis edwardii † Canis etruscus † Canis falconeri † Canis mosbachensis † Canis palmidens † Canis variabilis
150px |Cuon {{small|Hodgson, 1838}} | Cuon alpinus
150px |Lycaon {{small|Brookes, 1827}} | Lycaon pictus †Lycaon sekowei
150px |Lupulella {{small|Hilzheimer, 1906}} | Lupulella adusta Lupulella mesomelas
rowspan="5" style="text-align:center;"| Subtribe Cerdocyonina ! rowspan="5" style="text-align:center;"|The South American canines | 150px |Speothos {{small|Lund, 1839}} | Speothos venaticus †Speothos pacivorus
150px |Lycalopex {{small|Burmeister 1854}} | Lycalopex culpaeus Lycalopex fulvipes Lycalopex griseus Lycalopex gymnocercus Lycalopex sechurae Lycalopex vetulus †Lycalopex cultridens †Lycalopex ensenadensis
150px |Cerdocyon {{small|C. E. H. Smith, 1839}} | Cerdocyon thous
150px |Chrysocyon {{small|Smith, 1839}} | Chrysocyon brachyurus
150px |Atelocynus {{small|Cabrera, 1940}} | Atelocynus microtis
rowspan="3" style="text-align:center;"| Tribe Vulpini ! rowspan="3" style="text-align:center;"|The fox-like canines | 150px |Nyctereutes {{small|Temminck, 1838}} | Nyctereutes procyonoides Nyctereutes viverrinus †Nyctereutes abdeslami †Nyctereutes donnezani †Nyctereutes lockwoodi †Nyctereutes megamastoides †Nyctereutes sinensis †Nyctereutes tingi †Nyctereutes vinetorum
150px |Otocyon {{small|S. Müller, 1835}} | Otocyon megalotis
150px |Vulpes {{small|Garsault, 1764}} | Vulpes bengalensis Vulpes cana Vulpes chama Vulpes corsac Vulpes ferrilata Vulpes lagopus Vulpes macrotis Vulpes pallida Vulpes rueppellii Vulpes velox Vulpes vulpes Vulpes zerda †Vulpes hassani †Vulpes praeglacialis †Vulpes qiuzhudingi †Vulpes riffautae †Vulpes rooki †Vulpes skinneri †Vulpes stenognathus
rowspan="1" style="text-align:center;"| Genus Urocyon ! rowspan="1" style="text-align:center;"| Gray foxes | 150px |Urocyon {{small|Baird, 1857}} | U. cinereoargenteus U. littoralis † U. citrinus † U. galushai † U. minicephalus † U. progressus † U. webbi

DNA analysis shows that the first three form monophyletic clades. The wolf-like canines and the South American canines together form the tribe Canini.{{cite book |title=The Behavioural Biology of Dogs |last=Jensen |first=Per |year=2007 |publisher=CABI |isbn=978-1-84593-188-9 |pages=11–13 |url=https://books.google.com/books?id=SpkSd__EdKYC&pg=PA3 }} Molecular data imply a North American origin of living Canidae some 10 Mya and an African origin of wolf-like canines (Canis, Cuon, and Lycaon), with the jackals being the most basal of this group.

The South American clade is rooted by the maned wolf and bush dog, and the fox-like canines by the fennec fox and Blanford's fox. The gray fox and island fox are basal to the other clades; however, this topological difference is not strongly supported.{{cite journal |doi=10.1038/nature04338 |pmid=16341006 |title=Genome sequence, comparative analysis and haplotype structure of the domestic dog |journal=Nature |volume=438 |issue=7069 |pages=803–819 |year=2005 |last1=Lindblad-Toh |first1=Kerstin |last2=Wade |first2=Claire M. |last3=Mikkelsen |first3=Tarjei S. |last4=Karlsson |first4=Elinor K. |last5=Jaffe |first5=David B. |last6=Kamal |first6=Michael |last7=Clamp |first7=Michele |last8=Chang |first8=Jean L. |last9=Kulbokas |first9=Edward J. |last10=Zody |first10=Michael C. |last11=Mauceli |first11=Evan |last12=Xie |first12=Xiaohui |last13=Breen |first13=Matthew |last14=Wayne |first14=Robert K. |last15=Ostrander |first15=Elaine A. |last16=Ponting |first16=Chris P. |last17=Galibert |first17=Francis |last18=Smith |first18=Douglas R. |last19=Dejong |first19=Pieter J. |last20=Kirkness |first20=Ewen |last21=Alvarez |first21=Pablo |last22=Biagi |first22=Tara |last23=Brockmann |first23=William |last24=Butler |first24=Jonathan |last25=Chin |first25=Chee-Wye |last26=Cook |first26=April |last27=Cuff |first27=James |last28=Daly |first28=Mark J. |last29=Decaprio |first29=David |last30=Gnerre |first30=Sante |display-authors=6 |bibcode=2005Natur.438..803L |doi-access=free}}

The cladogram below is based on the phylogeny of Lindblad-Toh (2005) modified to incorporate recent findings on Canis, Vulpes,{{cite journal |last1=Zhao |first1=Chao |last2=Zhang |first2=Honghai |last3=Liu |first3=Guangshuai |last4=Yang |first4=Xiufeng |last5=Zhang |first5=Jin |title=The complete mitochondrial genome of the Tibetan fox (Vulpes ferrilata) and implications for the phylogeny of Canidae |journal=Comptes Rendus Biologies |volume=339 |issue=2 |year=2016 |pages=68–77 |issn=1631-0691 |doi=10.1016/j.crvi.2015.11.005 |pmid=26868757 |doi-access=free}} Lycalopex species,{{cite journal |first1=Ligia |last1=Tchaicka |first2=Thales Renato Ochotorena |last2=de Freitas |first3=Alex |last3=Bager |first4=Stela Luengos |last4=Vidal |first5=Mauro |last5=Lucherini |first6=Agustín |last6=Iriarte |first7=Andres |last7=Novaro |first8=Eli |last8=Geffen |first9=Fabricio Silva |last9=Garcez |first10=Warren E. |last10=Johnson |first11=Robert K. |last11=Wayne |first12=Eduardo |last12=Eizirik |display-authors=6 |year=2016 |title=Molecular assessment of the phylogeny and biogeography of a recently diversified endemic group of South American canids (Mammalia: Carnivora: Canidae) |journal=Genetics and Molecular Biology |volume=39 |issue=3 |pages=442–451 |doi=10.1590/1678-4685-GMB-2015-0189 |pmid=27560989 |pmc=5004827 |url=http://www.scielo.br/pdf/gmb/v39n3/1415-4757-gmb-1678-4685-GMB-2015-0189.pdf}} and Dusicyon.{{cite journal |author1=Slater, G. J. |author2=Thalmann, O. |author3=Leonard, J. A. |author4=Schweizer, R. M. |author5=Koepfli, K.-P. |author6=Pollinger, J. P. |author7=Rawlence, N. J. |author8=Austin, J. J. |author9=Cooper, A. |author10=Wayne, R. K. |display-authors=6 |year=2009 |title=Evolutionary history of the Falklands wolf |journal=Current Biology |volume=19 |issue=20 |pages=R937–R938 |doi=10.1016/j.cub.2009.09.018 |issn=0960-9822 |pmid=19889366 |bibcode=2009CBio...19.R937S |hdl=10261/58562 |s2cid=36185744 |hdl-access=free}}

{{Clade |style=font-size:80%; line-height:85%

|label1=Caninae

|1={{Clade

|1={{Clade

|label1=Canini

|1={{Clade

|style1=background-color:#ccccff;

|label1=Canina

|1={{Clade

|1={{Clade

|2=Lycaon pictus (African wild dog) 50 px

|1={{Clade

|2=Cuon alpinus (dhole) 50 px

|label1=Canis

|1={{Clade

|1={{Clade

|1={{Clade

|1={{Clade

|1={{Clade

|1=Canis latrans (coyote) 50 px

|2={{Clade

|1=Canis rufus (red wolf) 50px

|2=Canis lycaon (algonquin wolf) 50px

}}

}}

|2={{Clade

|1=Canis lupus (gray wolf) 50px

|2=Canis familiaris (domestic dog) 50px

}}

}}

|2=Canis anthus (African wolf) 50px

}}

|2=Canis simensis (Ethiopian wolf) 50px

}}

|2=Canis aureus (golden jackal) 50px

}}

}}

}}

|label2=Lupulella

|2={{Clade

|1=Lupulella adusta (side-striped jackal) 50 px

|2=Lupulella mesomelas (black-backed jackal) 50 px

}}

}}

|style2=background-color:#ccffcc;

|label2=Cerdocyonina

|2={{Clade

|1={{Clade

|1=Speothos venaticus (bush dog) 50 px

|2={{clade

|1=Chrysocyon brachyurus (maned wolf) 40 px

|2= †Dusicyon australis (Falkland Islands wolf)

}}

}}

|2={{Clade

|1={{Clade

|1={{Clade

|label1=Lycalopex

|1={{Clade

|1=Lycalopex vetulus (hoary fox) 40 px

|2={{Clade

|1=Lycalopex sechurae (Sechuran fox or Peruvian desert fox)

|2={{Clade

|1={{Clade

|1=Lycalopex fulvipes (Darwin's fox)

|2=Lycalopex gymnocercus (pampas fox) 50 px

}}

|2={{Clade

|1=Lycalopex griseus (South American gray fox or chilla)

|2=Lycalopex culpaeus (culpeo or Andean fox) 50 px

}}

}}

}}

}}

|2=Cerdocyon thous (crab-eating fox) 50 px

}}

|2=Atelocynus microtis (short-eared dog) 50 px

}}

}}

}}

}}

|style2=background-color:#ffcccc;

|label2=Vulpini

|2={{Clade

|1=Otocyon megalotis (bat-eared fox) 45 px

|2={{Clade

|1=Nyctereutes (raccoon dogs) 50 px

|label2=Vulpes

|2={{Clade

|1={{Clade

|1=Vulpes zerda (fennec fox) 50 px

|2=Vulpes cana (Blanford's fox) 50 px

}}

|2={{Clade

|1=Vulpes chama (Cape fox) 50 px

|2={{Clade

|1={{Clade

|1={{Clade

|1=Vulpes vulpes (red fox) 50 px

|2=Vulpes rueppellii (Ruppell's fox) 50 px

}}

|2={{clade

|1=Vulpes corsac (corsac fox) 50 px

|2=Vulpes ferrilata (Tibetan sand fox) 50 px

}}

}}

|2={{Clade

|1=Vulpes macrotis (kit fox) 50 px

|2=Vulpes lagopus (Arctic fox) 50 px

}}

}}

}}

}}

}}

}}

}}

|style2=background-color:#eeccFF;

|label2=Urocyon

|2={{Clade

|1=Urocyon littoralis (island fox) 50 px

|2=Urocyon cinereoargenteus (gray fox) 50 px

}}

}}

}}

{{reflist|refs=

{{cite journal |doi=10.1016/j.cub.2015.06.060 |pmid=26234211 |title=Genome-wide evidence reveals that African and Eurasian Golden Jackals are distinct species |journal=Current Biology |volume=25 |issue=16 |pages=2158–2165 |year=2015 |last1=Koepfli |first1=Klaus-Peter |last2=Pollinger |first2=John |last3=Godinho |first3=Raquel |last4=Robinson |first4=Jacqueline |last5=Lea |first5=Amanda |last6=Hendricks |first6=Sarah |last7=Schweizer |first7=Rena M. |last8=Thalmann |first8=Olaf |last9=Silva |first9=Pedro |last10=Fan |first10=Zhenxin |last11=Yurchenko |first11=Andrey A. |last12=Dobrynin |first12=Pavel |last13=Makunin |first13=Alexey |last14=Cahill |first14=James A. |last15=Shapiro |first15=Beth |last16=Álvares |first16=Francisco |last17=Brito |first17=José C. |last18=Geffen |first18=Eli |last19=Leonard |first19=Jennifer A. |last20=Helgen |first20=Kristofer M. |last21=Johnson |first21=Warren E. |last22=o'Brien |first22=Stephen J. |last23=Van Valkenburgh |first23=Blaire |last24=Wayne |first24=Robert K. |display-authors=6 |doi-access=free|bibcode=2015CBio...25.2158K }}{{rp|page=S1}}

{{cite book |last1=Miklosi |first1=Adam |title=Dog Behaviour, Evolution, and Cognition |publisher=Oxford University Press |edition=2 |series=Oxford Biology |year=2015 |pages=103–107 |url=https://books.google.com/books?id=VT-WBQAAQBAJ&pg=PA103 |via=Google Books |isbn=978-0199545667}}

{{cite journal |title=Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae) |year=2009 |first1=Richard |last1=Tedford |author1-link=Richard H. Tedford |first2=Xiaoming |last2=Wang |author2-link=Xiaoming Wang (paleontologist) |first3=Beryl E. |last3=Taylor |journal=Bulletin of the American Museum of Natural History |volume=325 |pages=1–218 |doi=10.1206/574.1 |hdl=2246/5999 |s2cid=83594819 |url=https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2009/issue-325/574.1/Phylogenetic-Systematics-of-the-North-American-Fossil-Caninae-Carnivora/10.1206/574.1.full}}

{{cite journal |last=Wayne |first=Robert K. |date=June 1993 |title=Molecular evolution of the dog family |journal=Trends in Genetics |volume=9 |issue=6 |pages=218–224 |url=http://wooferhouse.net/Links/MolecularEvolutionOfTheDogFamily/MolecularEvolutionOfTheDogFamily.htm |pmid=8337763 |doi=10.1016/0168-9525(93)90122-x}}

{{cite book |last1=Wang |first1=Xiaoming |author1-link=Xiaoming Wang (paleontologist) |last2=Tedford |first2=Richard H. |author2-link=Richard H. Tedford |title=Dogs: Their Fossil Relatives and Evolutionary History |publisher=Columbia University Press |location=New York |year=2008 |pages=1–232 |isbn=978-0-231-13529-0 |url={{Google books|plainurl=yes|id=LnWdpK7ctI0C|page=}} }}

}}

{{Carnivora|Ca.}}

{{Canidae extinct nav|C.}}

{{Taxonbar|from=Q2474088}}

Category:Extant Miocene first appearances