Phylogeny of the Caninae (Carnivora): Combining morphology, behaviour, genes and fossils

Abstract Phylogenetic relationships among 36 Recent and 42 extinct species of the Caninae (Canidae) were analysed, based on 360 morphological, developmental, ecological, behavioural and cytogenetic characters and 24 mitochondrial and nuclear markers. Primary phylogenetic analyses were accompanied by experimental analyses based on various combinations of data partitions and taxon samples. Leptocyon was recovered as a paraphyletic stem lineage of the Caninae; monophyly/paraphyly of the fox‐like canids (Vulpini) remains uncertain; Urocyon and Metalopex form a clade, possibly sister to all non‐ Leptocyon canids; Otocyon , Nyctereutes and Nurocyon form a clade; dog‐like canids (Canini) are monophyletic (with South American Cerdocyonina and Afro‐Holarctic Canina); all South American hypercarnivores ( Canis gezi , Protocyon , Speothos , Theriodictis ) form a clade, close to Chrysocyon and Dusicyon ; Canis arnensis , C. ferox , C. thooides , C. lepophagus and Eucyon spp. are basal to the Canina; Lycaon is an isolated African hypercarnivore; Cuon and its relatives ( Xenocyon , possibly also Canis antonii , C. falconeri and Cynotherium ) form a clade close to Canis s. str.; C. edwardii–C. etruscus–C. mosbachensis–C. palmidens–C. variabilis and hypercarnivorous Canis armbrusteri–C. dirus clades belong to Canis s. str. As the highly homoplastic morphological characters connected to dietary biology are the prominent characters available for the key fossil species, we conclude that macroevolutionary and palaeoecological analyses of the extinct and extant Caninae were to some extent compromised by the phylogenies used.