López-Torres and Silcox—European Paromomyidae
between Asia and Europe would have been rather difficult because the Turgai Strait would have posed a significant marine barrier to terrestrial mammals. The second wave of immigrant species (Phase 2) to Europe from North America was possible due to increased temperatures during the Paleocene-Eocene Thermal Maximum (PETM) that allowed high latitude dispersal. The second dispersal included the marsupial Peradectes Matthew and Granger, 1921 and the mesonychid Pachyaena Cope 1874 and is thought to have included paromomyids (Hooker, 2015; but see the following). European paromomyids were first reported from Pourcy,
France (Paleocene-EoceneMammal Zone [PE] III), by Louis and Michaux (1962) in a faunal list as an indeterminate species and genus of the ‘Phenacolemuridae,’ a family name now considered a junior synonym to Paromomyidae (Simpson, 1955). They were later ascribed to the North American paromomyid genus Phenacolemur Matthew, 1915 by Louis (1966), but no species level identification was given. Soon after, Russell et al. (1967) wrote the first comprehensive taxonomic study of European paromomyids in which they described two species from France: Phenacolemur fuscus from Mutigny, and Phenacolemur lapparenti fromAvenay (PE V). Later, Godinot (1984) described a new species, Arcius rougieri, fromPalette (PE II), France. In that paper, Godinot (1984) transferred the other two known species of European paromomyid into the newly named genus Arcius. Paromomyids have also been described from multiple early Eocene sites in France (Azillanet [Marandat, 1986], Condé- en-Brie [Louis, 1966;Aumont, 2003, 2004], Fordones [Marandat, 1991], Fournes [Marandat, 1991], Gland [Aumont, 2003], Grauves [Louis, 1970], Meudon [Russell et al., 1988, 1990], Prémontré [Dégremont et al., 1985], Rians [Godinot, 1981], Sézanne [Louis, 1970], Sotteville-sur-Mer [Smith et al., 2011], St. Agnan [Louis and Laurain, 1983], Venteuil [Aumont, 2003]), the United Kingdom (Abbey Wood [Hooker and Insole, 1980; Hooker, 1991, 1996, 1998]), Portugal (Silveirinha [Antunes, 1981; Estravís, 1992, 2000; Antunes et al., 1997]), and Spain (Masia de l’Hereuet [Marigó et al., 2012]). Godinot (1984) hypothesized that all paromomyids from Europe belong to a single genus, Arcius. A full analysis of the phylogenetic relationships of the
European paromomyids has never been published, although some ideas about possible relationships have been expressed. Russell et al. (1967) suggested that the European paromomyids were closely related to Phenacolemur; Godinot (1984) proposed A. rougieri as a potential ancestor to Arcius fuscus Russell et al., 1967 and Arcius lapparenti Russell et al., 1967 and that Arcius would be more closely related to Ignacius Matthew and Granger, 1921; Estravís (2000) advocated that A. zbyszewskii Estravís, 2000 was the most basal species of Arcius. The only cladistic analysis of the European paromomyids was included by Aumont (2003) in her unpublished dissertation, using 12 taxa and 35 dental characters. She found that A. fuscus and A. lapparenti formed a clade, with A. zbyszewskii as its sister taxon and A. rougieri being the most primitive lineage of the genus. The sister taxon for Arcius was the North American paromomyid Acidomomys hebeticus Bloch et al., 2002. This paper has three aims: (1) to revisit the alpha taxonomy
of the European Paromomyidae, (2) to analyze the phylogenetic relationships among European paromomyids and their rela- tionships to other members of the family using a data set that
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samples both characters and taxa more comprehensively than Aumont (2003), and (3) to discuss the biogeographic implica- tions of the phylogenetic analysis.
Repositories and institutional abbreviations.—British Museum of Natural History (BMNH), London, UK; Institut Català de Paleontologia Miquel Crusafont (IPS), Sabadell, Spain; MuséumNational d’Histoire Naturelle (Avenay collection, AV; Condé-en-Brie collection, CB; Mutigny collection, MU; Rians collection, RI), Paris, France; Royal Belgian Institute of Natural Sciences (Mammal collection, MAM), Brussels, Belgium; Universidade Nova de Lisboa (Silveirinha collection, SV), Lisbon, Portugal; Université de Montpellier (Fondones collec- tion, FDN; Fournes collection, FRN; Palette collection, PAT), Montpellier, France; University of Alberta Laboratory of Ver- tebrate Paleontology (UALVP), Edmonton, Canada; University of California Museum of Paleontology (UCMP), Berkeley, USA; University of Michigan Museum of Paleontology (UM), Ann Arbor, USA; United States Geological Survey collection (USGS), Washington, USA; United States National Museum (USNM) Washington, USA; Yale Peabody Museum (Princeton University collection, YPM-PU), New Haven, USA.
Systematic paleontology Order Primates Linnaeus, 1758
Superfamily Paromomyoidea (Simpson, 1940) Family Paromomyidae Simpson, 1940 Arcius Godinot, 1984
1962 ‘Phenacolemuridae’ gen. indet. Louis and Michaux, p. 171.
1966 Phenacolemur Louis, p. 51. 1967 Phenacolemur; Russell et al., p. 8, 12. 1970 Phenacolemur; Louis, p. 114. 1980 Phenacolemur; Hooker and Insole, p. 38. 1981 Phenacolemur; Antunes, p. 257. 1981 Phenacolemur; Antunes and Russell, p. 1101. 1981 Phenacolemur; Godinot, p. 77. 1983 Phenacolemur; Louis and Laurain, p. 9. 1984 Arcius Godinot, p. 85. 1985 Phenacolemur; Dégremont et al., p. 16. 1986 Arcius; Marandat, p. 88. 1988 Arcius; Russell et al., p. 432. 1991 Arcius; Marandat, p. 92. 1996 Arcius; Hooker, p. 209. 1998 Arcius; Hooker, p. 449. 2000 Arcius; Estravís, p. 283. 2003 Arcius; Aumont, pl. 1–24. 2004 Arcius; Aumont, fig. 3. 2010 Arcius; Hooker, p. 48. 2011 Arcius; Smith et al., fig. 19A, B. 2012 Arcius; Marigó et al., p. 430.
Type species.—Arcius rougieri.
Other species.—A. fuscus, A. lapparenti, A. zbyszewskii, Arcius hookeri n. sp., Arcius ilerdensis n. sp. (see descriptions below).
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