Project 46: R. V. Hill. 2005. Integration of Morphological Data Sets for Phylogenetic Analysis of Amniota: The Importance of Integumentary Characters and Increased Taxonomic Sampling. Systematic Biology. 54 (4):530-547.
Specimen: † Peltosaurus granulosus (AMNH:8138)
View: Lateral

Abstract

Several mutually exclusive hypotheses have been advanced to explain the phylogenetic position of turtles among amniotes. Traditional morphology-based analyses place turtles among extinct anapsids (reptiles with a solid skull roof), whereas more recent studies of both morphological and molecular data support an origin of turtles from within Diapsida (reptiles with a doubly fenestrated skull roof). Evaluation of these conflicting hypotheses has been hampered by nonoverlapping taxonomic samples and the exclusion of significant taxa from published analyses. Furthermore, although data from soft tissues and anatomical systems such as the integument may be particularly relevant to this problem, they are often excluded from large-scale analyses of morphological systematics. Here, conflicting hypotheses of turtle relationships are tested by (1) combining published data into a supermatrix of morphological characters to address issues of character conflict and missing data; (2) increasing taxonomic sampling by more than doubling the number of operational taxonomic units to test internal relationships within suprageneric ingroup taxa; and (3) increasing character sampling by approximately 25% by adding new data on the osteology and histology of the integument, an anatomical system that has been historically underrepresented in morphological systematics. The morphological data set assembled here represents the largest yet compiled for Amniota. Reevaluation of character data from prior studies of amniote phylogeny favors the hypothesis that turtles indeed have diapsid affinities. Addition of new ingroup taxa alone leads to a decrease in overall phylogenetic resolution, indicating that existing characters used for amniote phylogeny are insufficient to explain the evolution of more highly nested taxa. Incorporation of new data from the soft and osseous components of the integument, however, helps resolve relationships among both basal and highly nested amniote taxa. Analysis of a data set compiled from published sources and data original to this study supports monophyly of Amniota, Synapsida, Reptilia, Parareptilia, Eureptilia, Eosuchia, Diapsida, Neodiapsida, Sauria, Lepidosauria, and Archosauriformes, as well as several more highly nested divisions within the latter two clades. Turtles are here resolved as the sister taxon to a monophyletic Lepidosauria (squamates + Sphenodon), a novel phylogenetic position that nevertheless is consistent with recent molecular and morphological studies that have hypothesized diapsid affinities for this clade. [Amniota; integument; osteoderms; phylogeny; Testudines.]


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Project DOI: 10.7934/P46, http://dx.doi.org/10.7934/P46
This project containsMatrices
  • 753 Media
  • 1 Matrix
  • 222 Taxa
  • 428 Specimens
  • 368 Characters
Total size of project's media files: 699.16M

Download Project SDD File
Total scored cells: 17947
Total media associated with cells: 4454
Total labels associated with cell media: 0
Characters
Total characters: 368
Total characters with associated media: 0
Total characters with media with labels: 0
Total character states: 811
Total character states with associated media: 0
Total character states with media with labels:0
Total unordered/ordered characters:368/0
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MorphoBank Project 46

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    Authors' Institutions

    • Stony Brook University



    Members

    member name taxa specimens media media
    notes
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    notes  media
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    (scored, NPA, "-")
    cell
    notes  medialabels
    rules
    Seth Kaufman
    Project Administrator
    000036800017947
    (17947, 0, 0)
    0000
    MorphoBank Curator
    000000000
    (0, 0, 0)
    0000
    Robert Hill
    Full membership
    139376731000000
    (0, 0, 0)
    0000
    Mark Miller
    Full membership
    000000000
    (0, 0, 0)
    0000
    Maureen O'Leary
    Full membership
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    0000


    Taxonomic Overview for Matrix 'M147' (80 Taxa)

    taxon unscored
    cells
    scored
    cells
    no cell
    support
    NPA
    cells
    "-" cellscell images labels on
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    [1] † Seymouriidae
    Taxon name last Modified on 05/02/19
    1332351020015004
    [2] † Diadectomorpha
    Taxon name last Modified on 05/02/19
    151224770014304
    [3] † Casea
    Taxon name last Modified on 05/02/19
    14222622600004
    [4] † Ophiacodon retroversus
    Taxon name last Modified on 05/02/19
    137231850014604
    [5] † Edaphosauridae
    Taxon name last Modified on 05/02/19
    15321621500004
    [6] † Sphenacodontidae
    Taxon name last Modified on 05/02/19
    13223623600004
    [7] † Gorgonopsia
    Taxon name last Modified on 05/02/19
    14522322300004
    [8] † Procynosuchus
    Taxon name last Modified on 05/02/19
    13423623400004
    [9] † Morganucodon
    Taxon name last Modified on 05/02/19
    17519319300004
    [10] † Glyptodon reticulatus
    Taxon name last Modified on 05/02/19
    143226208002204
    [11] † Glyptotherium
    Taxon name last Modified on 05/02/19
    14822222000004
    [12] † Doedicurus giganteus
    Taxon name last Modified on 05/02/19
    144224207001704
    [13] † Panochthus frenzelianus
    Taxon name last Modified on 05/02/19
    143226208001704
    [14] Dasypus novemcinctus
    Taxon name last Modified on 12/05/13
    78295211008104
    [15] Zaedyus pichiy
    Taxon name last Modified on 12/05/13
    7829529000004
    [16] † Captorhinidae
    Taxon name last Modified on 05/02/19
    1312391180012104
    [17] † Araeoscelidia
    Taxon name last Modified on 05/02/19
    13423970023204
    [18] † Paleothyris acadiana
    Taxon name last Modified on 05/02/19
    15321521500004
    [19] † Claudiosaurus germaini
    Taxon name last Modified on 05/02/19
    18418418400004
    [20] † Younginiformes
    Taxon name last Modified on 05/02/19
    16521020300004
    [21] † Kuehneosauridae
    Taxon name last Modified on 05/02/19
    24812012000004
    [22] † Coahomasuchus kahleorum
    Taxon name last Modified on 05/02/19
    282868600004
    [23] † Desmatosuchus haplocerus
    Taxon name last Modified on 05/02/19
    148221153007804
    [24] † Typothorax coccinarum
    Taxon name last Modified on 05/02/19
    228140119002504
    [25] † Paratypothorax andressorum
    Taxon name last Modified on 05/02/19
    317515100004
    [26] † Pseudopalatus
    Taxon name last Modified on 05/02/19
    156214850013604
    [27] † Rutiodon
    Taxon name last Modified on 05/02/19
    128243191005304
    [28] † Protosuchus richardsoni
    Taxon name last Modified on 05/02/19
    142226169005804
    [29] † Akanthosuchus langstoni
    Taxon name last Modified on 05/02/19
    313585500004
    [30] † Simosuchus clarki
    Taxon name last Modified on 05/02/19
    172197470015104
    [31] † Goniopholis lucasii
    Taxon name last Modified on 05/02/19
    205163490011404
    [32] † Mahajangasuchus insignis
    Taxon name last Modified on 05/02/19
    164204260018204
    [33] Alligator mississippiensis
    Taxon name last Modified on 12/05/13
    153541350022904
    [34] Crocodylus
    Taxon name last Modified on 12/05/13
    173521940016104
    [35] Caiman crocodilus
    Taxon name last Modified on 12/05/13
    183502270012304
    [36] Gavialis gangeticus
    Taxon name last Modified on 12/05/13
    183502250012704
    [37] † Euoplocephalus tutus
    Taxon name last Modified on 05/02/19
    15521421300004
    [38] † Pinacosaurus grangeri
    Taxon name last Modified on 05/02/19
    166202760012904
    [39] † Sauropelta edwardsorum
    Taxon name last Modified on 05/02/19
    15421421400004
    [40] † Stegosaurus stenops
    Taxon name last Modified on 05/02/19
    12923923900004
    [41] † Scelidosaurus harrisonii
    Taxon name last Modified on 05/02/19
    20116816700004
    [42] † Scutellosaurus lawleri
    Taxon name last Modified on 05/02/19
    25811111000004
    [43] † Rapetosaurus krausei
    Taxon name last Modified on 05/02/19
    342268001804
    [44] † Saltasaurus loricatus
    Taxon name last Modified on 05/02/19
    286838200004
    [45] † Titanosauridae sp
    Taxon name last Modified on 05/02/19
    343252500004
    [46] † Proganochelys quenstedti
    Taxon name last Modified on 05/02/19
    141227550017304
    [47] Chelydra serpentina
    Taxon name last Modified on 12/05/13
    643041950011004
    [48] Pseudemys scripta
    Taxon name last Modified on 12/05/13
    68300208009504
    [49] Geochelone
    Taxon name last Modified on 12/05/13
    693011780012404
    [50] Trionyx
    Taxon name last Modified on 12/05/13
    7429529400004
    [51] † Baena
    Taxon name last Modified on 05/02/19
    2709832007004
    [52] Sphenodon punctatus
    Taxon name last Modified on 12/05/13
    553131200019604
    [53] Heloderma
    Taxon name last Modified on 12/05/13
    19350339001004
    [54] Corucia zebrata
    Taxon name last Modified on 12/05/13
    1535335300004
    [55] Tiliqua
    Taxon name last Modified on 12/05/13
    1735335100004
    [56] Cordylus
    Taxon name last Modified on 12/05/13
    153551250023204
    [57] Zonosaurus
    Taxon name last Modified on 12/05/13
    163551790017804
    [58] Anguis fragilis
    Taxon name last Modified on 12/05/13
    812871680012004
    [59] Ophisaurus
    Taxon name last Modified on 12/05/13
    7529429300004
    [60] Gerrhonotus
    Taxon name last Modified on 12/05/13
    1135835700004
    [61] Elgaria
    Taxon name last Modified on 12/05/13
    1535335300004
    [62] † Glyptosaurinae
    Taxon name last Modified on 05/02/19
    221147370011304
    [63] † Mesosauridae
    Taxon name last Modified on 05/02/19
    19916916900004
    [64] † Millerettidae
    Taxon name last Modified on 05/02/19
    16021220800004
    [65] † Anthodon serrarius
    Taxon name last Modified on 05/02/19
    12024824800004
    [66] † Scutosaurus karpinskii
    Taxon name last Modified on 05/02/19
    9927126900004
    [67] † Bradysaurus seeleyi
    Taxon name last Modified on 05/02/19
    10226626600004
    [68] † Pareiasaurus serridens
    Taxon name last Modified on 05/02/19
    11025825800004
    [69] † Procolophon
    Taxon name last Modified on 05/02/19
    137234840014904
    [70] † Owenetta rubidgei
    Taxon name last Modified on 05/02/19
    19817117000004
    [71] † Placodus
    Taxon name last Modified on 05/02/19
    1232451350011304
    [72] † Eosauropterygia
    Taxon name last Modified on 05/02/19
    22317214500004
    [73] † Cyamodontoidea
    Taxon name last Modified on 05/02/19
    165212670013904
    [74] † Rhynchosauria
    Taxon name last Modified on 05/02/19
    151226980011904
    [75] † Trilophosaurus buettneri
    Taxon name last Modified on 05/02/19
    16820120000004
    [76] † Choristodera
    Taxon name last Modified on 05/02/19
    16620320200004
    [77] † Prolacertiformes
    Taxon name last Modified on 05/02/19
    15422621400004
    [78] † Acleistorhinus pteroticus
    Taxon name last Modified on 05/02/19
    25411411400004
    [79] † Lanthanosuchidae
    Taxon name last Modified on 05/02/19
    25112011700004
    [80] † Macroleter poezicus
    Taxon name last Modified on 05/02/19
    21815015000004


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