The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis

  • Timothy J. GaudinEmail author
  • Robert J. Emry
  • John R. Wible
Original Paper


The present study was undertaken in order to effect a comprehensive phylogenetic analysis of the order Pholidota, examining seven of the eight currently recognized extant species (absent is Manis culionensis, formerly recognized as a subspecies of Manis javanica) and nearly all the well-known fossil taxa, and employing a wide range of osteological characters from the entire skeleton. In addition, the relationship of pangolins to several putative early Tertiary relatives, including palaeanodonts and the enigmatic “edentate” Eurotamandua joresi, were investigated. The goal of the study was to improve understanding of the systematics and the biogeographic and evolutionary history of the pangolins. A computer-based cladistic analysis of phylogenetic relationships among seven extant species of pangolins, five extinct pangolin species (including all but one of the well-preserved taxa), as well as Eurotamandua and two genera of metacheiromyid palaeanodonts, Palaeanodon and Metacheiromys, was performed based upon 395 osteological characteristics of the skull and postcranial skeleton. Characters were polarized via comparison to the following successive outgroups: the basal feliform carnivoran Nandinia binotata and the hedgehog Erinaceus sp., a eulipotyphlan laursiatherian placental. A revised classification is presented based on the results of the analysis. The results support the monophyly of Pholidota and Palaeanodonta by providing new anatomical characters that can serve to diagnose a pangolin/palaeanodont clade, termed here Pholidotamorpha. Pholidota is defined so as to include all living and fossil pangolins, including all three taxa of middle Eocene “edentates” from the Messel fauna of Germany, among them Eurotamandua joresi. The results do not support the monophyly of the remaining two Messel “edentates” originally placed in the same genus Eomanis, which is restricted to the type species Eomanis waldi. Euromanis, new genus, is named with Eomanis krebsi Storch and Martin, 1994, as the type species, to form a new combination Euromanis krebsi (Storch and Martin, 1994). The analysis strongly supports the monophyly of a crown clade of pangolins diagnosed by many anatomical synapomorphies, the family Manidae. This crown clade is sister to the family Patriomanidae, which includes two Tertiary taxa, Patriomanis americana and Cryptomanis gobiensis, within the superfamily Manoidea. The relationship of the Tertiary European pangolin Necromanis to these two families is unresolved. Within Manidae, the extant species are divided into three well-supported, monophyletic genera, Manis for the Asian pangolins, Smutsia for the African ground pangolins, and Phataginus for the African tree pangolins. The latter two form a monophyletic African assemblage, the subfamily Smutsiinae. The biogeographic implications of this phylogeny are examined. A European origin for Pholidota is strongly indicated. The fossil record of pangolins would seem to support a European origin for the modern forms, with subsequent dispersal into sub-Saharan African and then to southern Asia, and the phylogeny produced in this analysis is consistent with such a scenario.


Morphology Pholidota Palaeanodonta Pangolins Phylogeny Eurotamandua Euromanis krebsi 



We wish to thank the following institutions and individuals for providing access to the specimens that formed the basis of this study: Ross MacPhee, Richard Monk, Nancy Simmons, and Eileen Westwig, Department of Mammalogy, American Museum of Natural History, New York, NY; John Flynn and Meng Jin, Department of Vertebrate Paleontology, American Museum of Natural History, New York, NY; Bruce Patterson, Larry Heaney, and Bill Stanley, Division of Mammals, Field Museum of Natural History, Chicago, IL; Prof. Gerhard Storch, Forschungs-Institut Senckenberg, Frankfurt am Main, Germany; Norbert Micklich and Connie Kurz, Hessisches Landesmuseum Darmstadt, Darmstadt, Germany; Ken Rose, Johns Hopkins University, Baltimore, MD; Richard, Thorington, Linda Gordon, and Helen Kafka, Department of Mammalogy, National Museum of Natural History, Washington, D.C.; and Eberhard “Dino” Frey, Staatliches Museum für Naturkunde, Karlsruhe, Germany. For help with the preparation and casting of fossil specimens connected with this study, we thank Fred Grady, Pete Kroehler, and Steve Jabo of the Department of Paleobiology at the National Museum of Natural History. We gratefully acknowledge Jeremy Jacobs of the Department of Mammalogy, National Museum of Natural History, who prepared radiographs from alcohol specimens of extant pangolins that were useful for coding portions of the matrix. Special thanks go to Prof. Gerhard Storch, who provided access to new, undescribed material of Necromanis for the purposes of coding this taxon, and who also provided us casts of several specimens of Eomanis waldi and the type of Euromanis krebsi, and showed us his three-dimensional x-ray photographs of Eurotamandua joresi. His material help and insightful comments were critical to the success of this project. We thank Ken Rose (Johns Hopkins University); Jeremy Bramblett, Stylianos Chatzimanolis, Charles Nelson, and Joey Shaw (University of Tennessee at Chattanooga); and John Rawlins (Carnegie Museum of Natural History) for useful discussions concerning this study; and, Michelle Spaulding (American Museum of Natural History) for providing information on Nandinia. In addition, this manuscript was greatly improved by the comments of three anonymous reviewers. We thank Julia Morgan Scott for her skillful work in the preparation of illustrations. For housing T. Gaudin during his 2002 period of sabbatical study, and once again during the summer of 2004, special thanks goes to the Department of Paleobiology at the National Museum of Natural History. T. Gaudin’s work on this project was supported by a sabbatical grant from the University of Chattanooga Foundation, and by NSF RUI Grant DEB 0107922 and NSF AToL Grant 0629959; J.R. Wible was supported by NSF AToL Grant 0629959.


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Copyright information

© Springer Science+Business Media, LLC 2009

Authors and Affiliations

  • Timothy J. Gaudin
    • 1
    Email author
  • Robert J. Emry
    • 2
  • John R. Wible
    • 3
  1. 1.Department of Biological & Environmental SciencesUniversity of Tennessee at ChattanoogaChattanoogaUSA
  2. 2.Department of Paleobiology, MRC 121, National Museum of Natural HistorySmithsonian InstitutionWashingtonUSA
  3. 3.Section of MammalsCarnegie Museum of Natural HistoryPittsburghUSA

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