López-Torres and Silcox—European Paromomyidae
933
alveoli for that tooth are not preserved. It is, therefore, impos- sible to determine how many roots would have been present. In light of these considerations, we would argue that there is no support for an Acidomomys-Arcius clade, which Aumont (2003) tentatively named as the tribe Arciini (incorrectly reported by Aumont [2003, p. 364] as ‘Arciusini’). In addition, the fact that Acidomomys appears as the sister taxon to Arcius might explain why Aumont’s phylogeny differs from those reported here in that Arcius rougieri represents the most primitive lineage of European paromomyids in her analysis, instead of Arcius zbyszewskii as found here. When Arcius hookeri n. sp. is included in the analysis
(Analysis 2; Fig. 10.2), it also yields one tree unequivocally. Arcius hookeri is placed as the sister taxon of a clade that includes A. fuscus, A. lapparenti, and A. rougieri. This result supports the inference that the Arcius specimen from England pertains to a separate lineage and therefore should be named as a distinct species. The clade that includes A. hookeri, A. fuscus, A. lapparenti, and A. rougieri is supported by the loss of the paraconid on the M3, which is clearly present in the more primitive A. zbyszewskii. The addition of the paromomyid specimen from Sotteville-
Figure 11. Cladogram generated by strict consensus of six trees. All European paromomyid taxa were included in this tree.
primitive lineage of European paromomyids, opening up the possibility of it being an ancestral species to all European par- omomyids. In terms of the rest of paromomyid relationships, all of the trees from this paper agree with Bloch et al. (2002) and Aumont (2003) that Phenacolemur and Ignacius are closely related and that Paromomys is a very primitive member of the family. However, the position of Acidomomys seems more contentious. Whereas Acidomomys makes a clade with Ignacius in our trees, Bloch et al. (2002) found Acidomomys to be the sister taxon of the clade Ignacius+ Phenacolemur, and Aumont’s (2003) analysis resulted in Acidomomys being the sister taxon to Arcius. Aumont (2003) stated that the two char- acters that support the Acidomomys-Arcius clade are “l’absence de la p3” (absence of P3) and “la presence d’une P3 uniradiculée” (presence of a single-rooted P3) (Aumont, 2003, p. 364). However, Acidomomys hebeticus does have a P3 (Bloch et al., 2002), and therefore that trait seems to be incorrectly coded in Aumont’s (2003) matrix. In addition, the other char- acter (presence of a single-rooted P3) seems to be misreported because P3 is coded in the matrix as being double rooted for both Acidomomys hebeticus and Arcius rougieri and not observable (i.e., question mark) in the rest of Arcius species. While it is clear that Acidomomys hebeticus had a double-rooted P3 (Bloch et al., 2002), as coded, the number of roots for P3 in Arcius is difficult to assess. Only a small piece of the distobuccal aspect of the P3 of A. rougieri is preserved (Godinot, 1984), and the
sur-Mer (Normandy, France) also results in a single most parsimonious tree (Analysis 3; Fig. 10.3). In this case, the Normandy specimen (MAM 6 STV 2008) forms a clade with Arcius zbyszewskii. This result is consistent with the morpho- logical resemblance between the fossils from Portugal and the one from Normandy and supports the inference that all these specimens belong to the same species, A. zbyszewskii. When including Arcius ilerdensis n. sp. in the analysis
(Analysis 4), the clade A. fuscus-A. lapparenti-A. rougieri, present in the results from Analyses 1–3, collapses (Fig. 10.4). In three of the six trees recovered from this analysis (Fig. S1.1, S1.2, S1.6), A. ilerdensis falls out as being most closely related to A. rougieri. Arcius ilerdensis also appears in two other trees (Fig. S1.4, S1.5) as the sister group to the clade A. fuscus- A. lapparenti-A. rougieri, and as most closely related to A. lapparenti in another tree (Fig. S1.3). When all species are included in the analysis (Analysis 5;
Fig. 11), most Arcius lineages collapse, likely because of the large amounts of missing data for several taxa. However, it is important to note that the clade including A. fuscus and A. lapparenti still holds in this cladogram. In terms of the other genera of paromomyids, our results
show that the sister group of Arcius is not Acidomomys,as reported in Aumont (2003), but the clade that includes Phenacolemur, Ignacius, and Acidomomys. The genus Acidomomys is most closely related to Ignacius. Paromomys and Edworthia appear as members of the most primitive lineage of paromomyids. These results also support the idea from Gunnell (1989) that the family Paromomyidae could be subdivided into two subfamilies: the Paromomyinae and the Phenacolemurinae. According to Gunnell (1989), based on the genera described at that time, paromomyines would only be composed of the genus Paromomys, and phenacolemurines would include Phenacolemur, Ignacius, and Elwynella. Here, Paromomyinae would be extended to include Paromomys and Edworthia, and Phenacolemurinae would include Phenacolemur, Ignacius, Arcius, and possibly Elwynella.
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