Evolution of Lagomorphs luis a. ruedas, josé m. mora, and hayley c. lanier What, if anything, is a rabbit? (Wood 1957:417) As currently understood, the order Lagomorpha has two living families: the Ochotonidae (pikas) and the Leporidae (cottontails and hares or jackrabbits). There are 29 extant species of pika in the genus Ochotona, whereas the Leporidae includes 63 species in 11 genera. The Sardinian pika, †Prolagus sardus, represents a third, recently extinct family of pika relatives, the †Prolagidae. However, and notwithstanding this apparent simplicity, both the taxonomic and the evolutionary histories of the Lagomorpha are complex and have been extensively debated. The original taxonomic understanding of the Lagomorpha (Linnaeus 1758) was unusual by modern standards. Linnaeus recognized eight orders of mammals, ranging taxonomically from what we might today consider a family (the whales, “Cetae”) to strange collections of subclasses (his “Bestiæ” and “Glires”). Lagomorphs were in the order Glires, which included six genera: Rhinoceros (rhinoceroses, two species), Hystrix (porcupines, five species), Lepus (hares and cottontails, four species), Castor (beaver, two species), Mus (mice and allies, 15 species), and Sciurus (squirrels, seven species). Thus the first taxonomic association of Lagomorpha was in what we might today consider a group of several distinct orders including rodents and, incongruously, an odd- toed ungulate: the rhinoceros. Linnaeus’s “genera” in Glires are in fact akin to modern families, if not orders. Linnaeus did not include pikas in the definitive tenth edition of his nomenclatural treatise; the four species included in Lepus were L. timidus, L. capensis, “Lepus” (= Oryctolagus) cuniculus, and “Lepus” (= Sylvilagus) brasiliensis. As a bit of trivia, the legendary modern “jackalope” appears to have been known to Linnaeus, who noted (1758:57) that the horned hare, if not a hybrid, was almost certainly mythical. In the particular instance of Lepus in the tenth edition, Linnaeus’s family concept was identical to what today we would call Leporidae. However, his diagnosis of the genus Lepus based on the duplicated first pair of upper incisors is in fact the modern diagnosis of the order Lagomorpha (Gidley 1912). Illiger (1811) used that character to erect the family Duplicidentata (still within the rodents), including two genera: Lepus and Lagomys (= Ochotona). Fischer called this same family- level group of Lepus and Lagomys the Leporini (Fischer 1817:372) and Leporinorum (Fischer 1817:409). The first use of “Leporidae” was by Gray (1821:304); however, he used the term in the same sense as Illiger and Fischer, to include Lepus and Lagomys (thus corresponding to Lagomorpha) in his order Rosores. Waterhouse (1839) later split “rodents” into three higher categories, or sections, the third of which, Leporina, corresponded to the contemporary concept of Lagomorpha. It was Brandt (1855:295) who eventually coined for the group the now commonly used term “Lagomorpha” (“hare shaped”), also at the subordinal rank within rodents (along with Sciuromorpha, Myomorpha, and Hystricomorpha). In addition, although he clearly called lagomorphs a suborder (“Subordo IV. Lagomorphi seu 1709048_int_cc2015.indd 4 15/9/2017 15:59 Evolution of Lagomorph 5 Lagomorpha”), Brandt (1855:295) began his discussion of the groups with the words “Ordo Leporinus.,” underscoring their stark distinction from rodents, based on the presence of four upper incisors in lagomorphs. Thus, while it could be argued that Brandt was the first to suggest ordinal status for lagomorphs, it was not until 1912 that J. W. Gidley formally called for ordinal rank for the Lagomorpha. Evolutionary History and Chronology of the Lagomorpha—Interordinal Relationships The early evolutionary diversification of Glires, the group including both rodents and lagomorphs, has been hypothesized to have occurred in the Late Cretaceous, immediately preceding the Cretaceous- Paleogene boundary (Asher et al. 2005), some 66 mya (near the time that non- avian dinosaurs went extinct). Evidence for this early diversification comes from the extinct †Mimotonidae, a non- natural grouping (in this instance, not including all descendants from a last common ancestor) of Glires forming with Lagomorpha the Duplicidentata, and the acknowledged closest relatives to Lagomorpha (Asher et al. 2005). Successive sister groups included †Gomphos elkema and also the non- natural †Eurymylidae (Asher et al. 2005). Within Lagomorpha, as currently understood, the earliest known fossil dates to at least 53 mya and is known from Early Eocene Cambay Shale deposits in west- central India from the Vastan lignite mine (Rose et al. 2008). Prior to that report, the oldest Lagomorpha were dated to the Irdinmanhan (48.6–37.2 mya; Middle Eocene) of Asia.