Project 291: T. J. Gaudin, R. J. Emry, J. R. Wible. 2009. The phylogeny of living and extinct pangolins (Mammalia, Pholidota) and associated taxa: a morphology based analysis. Journal of Mammalian Evolution. 16 (4):235-305.
Specimen: Eomanis waldi (SMF MEA:263)

Abstract

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 Superfamily Manoidea. This crown clade can in turn be divided into two monophyletic families: the Patriomanidae, which includes two Tertiary taxa, Patriomanis americana and Cryptomanis gobiensis, and the Manidae, which includes all the extant pangolins. 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 scenario.


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Project DOI: 10.7934/P291, http://dx.doi.org/10.7934/P291
This project containsMatrices
  • 63 Media
  • 1 Matrix
  • 1 Document
  • 21 Taxa
  • 18 Specimens
  • 395 Characters
Total size of project's media files: 3.2M

Download Project SDD File
Total scored cells: 5602
Total media associated with cells: 0
Total labels associated with cell media: 0
Characters
Total characters: 395
Total characters with associated media: 0
Total characters with media with labels: 0
Total character states: 1013
Total character states with associated media: 113
Total character states with media with labels:113
Total unordered/ordered characters:395/0
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MorphoBank Project 291

    This research
    supported by

    Authors' Institutions

    • University of Tennessee

    • Carnegie Museum of Natural History

    • Smithsonian Institution



    Members

    member name taxa specimens media chars character
    media
    labels
    cell scorings
    (scored, NPA, "-")
    cell
    notes  medialabels
    rules
    Timothy Gaudin
    Project Administrator
    2118633951161215602
    (5602, 0, 0)
    0000
    MorphoBank Curator
    Full membership
    0000000
    (0, 0, 0)
    0000
    Robert Emry
    Full membership
    0000000
    (0, 0, 0)
    0000
    John Wible
    Full membership
    0000000
    (0, 0, 0)
    0000


    Taxonomic Overview for Matrix 'M731' (17 Taxa)

    taxon unscored
    cells
    scored
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    no cell
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    NPA
    cells
    "-" cellscell images labels on
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    [1] Erinaceus sp.
    Last Modified in 12/05/13
    2836936700004
    [2] Nandinia binotata
    Last Modified in 12/05/13
    1138638400004
    [3] Manis crassicaudata
    Last Modified in 12/05/13
    1338938200004
    [4] Manis javanica
    Last Modified in 12/05/13
    143639400004
    [5] Manis pentadactyla
    Last Modified in 12/05/13
    641738900004
    [6] Smutsia gigantea
    Last Modified in 12/05/13
    545039000004
    [7] Smutsia temminckii
    Last Modified in 12/05/13
    840538700004
    [8] Phataginus tetradactyla
    Last Modified in 12/05/13
    141739400004
    [9] Phataginus tricuspis
    Last Modified in 12/05/13
    044539500004
    [10] Patriomanis americana
    Last Modified in 12/05/13
    3536936000004
    [11] Cryptomanis gobiensis
    Last Modified in 12/05/13
    17222522300004
    [12] Necromanis sp.
    Last Modified in 12/05/13
    19520520000004
    [13] Eomanis waldi
    Last Modified in 12/05/13
    15324824200004
    [14] Euromanis krebsi
    Last Modified in 12/05/13
    309898600004
    [15] Eurotamandua joresi
    Last Modified in 12/05/13
    17922121600004
    [16] Palaeanodon sp.
    Last Modified in 12/05/13
    18621420900004
    [17] Metacheiromys sp.
    Last Modified in 12/05/13
    7931731600004


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    Specimen list13425
    Taxon list8147
    Bibliography976
    Documents list1138




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