Abstract
The Aplodontoidea, now restricted to only one North American species (Aplodontia rufa), have shown a wide Holarctic extension since the Upper Eocene. As their fossil record is poor, their phylogenetic relationships and the origins of their successive radiations remain unclear. We perform here phylogenetic analyses, primarily based on dental evidence (94 dental of 97 characters), restricted to Paleogene and early Miocene taxa (46 taxa) in order to avoid biases introduced by substantially derived (divergent) taxa. We confirm the inclusion of some problematic genera such as Cedromus or Douglassciurus within Sciuroidea rather than in Aplodontoidea. Ephemeromys and Lophallomys appear as the most basal members of the Aplodontoidea, and Epeiromys is the closest outgroup of the Sciuroidea-Aplodontoidea clade. The relationships among the “prosciurines” remain unclear, with paraphyletic genera such as Prosciurus and Haplomys. Their diagnoses are reevaluated and a new genus is described. The Aplodontidae, including the clade of the latter, and Haplomys liolophus display a dichotomy between Ansomyinae and Aplodontinae, the two crown groups. The first clade formed by the European species argoviensis and descendens (referred to a new genus) can be proposed as a sister group of the species of Ansomys. The second branch of the dichotomy includes the European Plesispermophilus and Sciurodon as basal groups. The species of Parallomys do not form a clade, and the genus appears paraphyletic. The last dichotomy separates the Allomys clade from the ‘meniscomyine’ clade. Comparisons of the selected species allow consideration of their patterns of dental evolution (e.g. enlargement of P4, development of a metaloph—protoloph disto-mesial connection, of crescentic shape in main cusps and ectoloph, of a buccal protruding compressed mesostyle, of a metastylid crest or an anterior spur of the hypoconid, etc.). The split between sciuroids and aplodontoids occurred in North America, and then aplodontoids dispersed rapidly throughout the whole Holarctic region. The first aplodontid adaptive radiation took place either in North America or in Asia. Periodic exchanges occurred between Europe, Asia and North America, and the last radiations (meniscomyines) were restricted in North America.
Kurzfassung
Die Aplodontoidea waren seit dem Ober-Eozän holarktisch weit verbreitet, sind heute aber auf eine nordamerikanische Art (Aplodontia rufa) beschränkt. Da ihre fossilen Belege spärlich sind, bleiben ihre phylogenetischen Beziehungen sowie die Wurzeln ihrer aufeinander folgenden Radiationen unklar. Wir haben eine phylogenetische Analyse der Aplodontoidea durchgeführt, die in erster Linie auf Zahnmerkmalen basiert (94 von 97 Merkmalen). Die Auswahl der Taxa ist auf das Paläogen und frühe Miozän beschränkt (46 Taxa), um eine Verzerrung durch stark abgeleitete (divergente) Taxa zu vermeiden. Nach unserer Analyse gehören einige bisher fragliche Sciuroidea wie Cedromus oder Douglassciurus tatsächlich in diese Gruppe und nicht zu den Aplodontoidea. Ephemeromys und Lophallomys erweisen sich als ursprünglichste Mitglieder der Aplodontoidea, und Epeiromys stellt die Schwestergruppe des von Sciuroidea und Aplodontoidea gebildeten Taxons dar. Die Beziehungen zwischen den “Prosciurinen” bleiben unklar mit paraphyletischen Gattungen wie Prosciurus und Haplomys. Ihre Diagnosen werden revidiert und eine neue Gattung wird beschrieben. Die Aplodontidae inklusive der neuen Gattung und von Haplomys liolophus bilden eine Dichotomie mit Ansomyinae und Aplodontinae. Der erste Zweig, der aus den europäischen Arten argoviensis und descendens (zu einer neuen Gattung gestellt) besteht, bildet die Schwestergruppe zur Gattung Ansomys. Der zweite Ast der Dichotomie umfasst den europäischen Plesispermophilus und auch Sciurodon als basale Gruppen. Die Arten von Parallomys stellen kein Monophylum dar, und die Gattung erscheint paraphyletisch. Die letzte Dichotomie trennt den Allomys-Zweig von dem ‘meniscomyinen’ Zweig. Ein Vergleich der ausgewählten Arten zeigt die evolutiven Veränderungen im Gebiss (z.B. Vergrößerung des P4, Entwicklung einer disto-mesialen Verbindung zwischen Metaloph und Protoloph, eines bogenförmigen Verlaufs von Haupthöckern und Ectoloph, eines Metastylid-Grates oder eines anterioren Vorsprungs des Hypoconids usw….). Die Aufspaltung von Sciuroiden und Aplodontoiden erfolgte in Nordamerika, und darauf verbreiteten sich die Aplodontoiden im gesamten holarktischen Bereich. Die erste adaptive Radiation der Aplodontoiden erfolgte entweder in Nordamerika oder in Asien. Es kam zu wiederholtem Austausch zwischen Europa, Asien und Nordamerika, aber die letzten Radiationen (Meniscomyinen) waren auf Nordamerika beschränkt.
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Acknowledgments
This paper has been supported by the “Institut des Sciences de l’Evolution”, UMR5554 CNRS—Université Montpellier 2 and L.M. by ANR-08-JCJC-0017 (PALASIAFRICA). The Ulantatal specimens are the result of the Chinese/German expedition in 1987, with the support of the Academia Sinica (China), the Max Planck-Gesellschaft (Germany) and the Bureau of Geology and Mineral Resources of Ningxia Hui Autonomous Region (China). Publication ISE-M no 2012-085.
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Vianey-Liaud, M., Gomes Rodrigues, H. & Marivaux, L. Early adaptive radiations of Aplodontoidea (Rodentia, Mammalia) on the Holarctic region: systematics, and phylogenetic and paleobiogeographic implications. Paläontol Z 87, 83–120 (2013). https://doi.org/10.1007/s12542-012-0143-3
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DOI: https://doi.org/10.1007/s12542-012-0143-3