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



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    Seth Kaufman
    Project Administrator
    000036800017947
    (17947, 0, 0)
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    MorphoBank Curator
    000000000
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    Robert Hill
    Full membership
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    (0, 0, 0)
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    Mark Miller
    Full membership
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    Maureen O'Leary
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    Taxonomic Overview for Matrix 'M147' (80 Taxa)

    taxon unscored
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    [1] † Seymouriidae
    Last Modified in 05/02/19
    1332351020015004
    [2] † Diadectomorpha
    Last Modified in 05/02/19
    151224770014304
    [3] † Casea
    Last Modified in 05/02/19
    14222622600004
    [4] † Ophiacodon retroversus
    Last Modified in 05/02/19
    137231850014604
    [5] † Edaphosauridae
    Last Modified in 05/02/19
    15321621500004
    [6] † Sphenacodontidae
    Last Modified in 05/02/19
    13223623600004
    [7] † Gorgonopsia
    Last Modified in 05/02/19
    14522322300004
    [8] † Procynosuchus
    Last Modified in 05/02/19
    13423623400004
    [9] † Morganucodon
    Last Modified in 05/02/19
    17519319300004
    [10] † Glyptodon reticulatus
    Last Modified in 05/02/19
    143226208002204
    [11] † Glyptotherium
    Last Modified in 05/02/19
    14822222000004
    [12] † Doedicurus giganteus
    Last Modified in 05/02/19
    144224207001704
    [13] † Panochthus frenzelianus
    Last Modified in 05/02/19
    143226208001704
    [14] Dasypus novemcinctus
    Last Modified in 12/05/13
    78295211008104
    [15] Zaedyus pichiy
    Last Modified in 12/05/13
    7829529000004
    [16] † Captorhinidae
    Last Modified in 05/02/19
    1312391180012104
    [17] † Araeoscelidia
    Last Modified in 05/02/19
    13423970023204
    [18] † Paleothyris acadiana
    Last Modified in 05/02/19
    15321521500004
    [19] † Claudiosaurus germaini
    Last Modified in 05/02/19
    18418418400004
    [20] † Younginiformes
    Last Modified in 05/02/19
    16521020300004
    [21] † Kuehneosauridae
    Last Modified in 05/02/19
    24812012000004
    [22] † Coahomasuchus kahleorum
    Last Modified in 05/02/19
    282868600004
    [23] † Desmatosuchus haplocerus
    Last Modified in 05/02/19
    148221153007804
    [24] † Typothorax coccinarum
    Last Modified in 05/02/19
    228140119002504
    [25] † Paratypothorax andressorum
    Last Modified in 05/02/19
    317515100004
    [26] † Pseudopalatus
    Last Modified in 05/02/19
    156214850013604
    [27] † Rutiodon
    Last Modified in 05/02/19
    128243191005304
    [28] † Protosuchus richardsoni
    Last Modified in 05/02/19
    142226169005804
    [29] † Akanthosuchus langstoni
    Last Modified in 05/02/19
    313585500004
    [30] † Simosuchus clarki
    Last Modified in 05/02/19
    172197470015104
    [31] † Goniopholis lucasii
    Last Modified in 05/02/19
    205163490011404
    [32] † Mahajangasuchus insignis
    Last Modified in 05/02/19
    164204260018204
    [33] Alligator mississippiensis
    Last Modified in 12/05/13
    153541350022904
    [34] Crocodylus
    Last Modified in 12/05/13
    173521940016104
    [35] Caiman crocodilus
    Last Modified in 12/05/13
    183502270012304
    [36] Gavialis gangeticus
    Last Modified in 12/05/13
    183502250012704
    [37] † Euoplocephalus tutus
    Last Modified in 05/02/19
    15521421300004
    [38] † Pinacosaurus grangeri
    Last Modified in 05/02/19
    166202760012904
    [39] † Sauropelta edwardsorum
    Last Modified in 05/02/19
    15421421400004
    [40] † Stegosaurus stenops
    Last Modified in 05/02/19
    12923923900004
    [41] † Scelidosaurus harrisonii
    Last Modified in 05/02/19
    20116816700004
    [42] † Scutellosaurus lawleri
    Last Modified in 05/02/19
    25811111000004
    [43] † Rapetosaurus krausei
    Last Modified in 05/02/19
    342268001804
    [44] † Saltasaurus loricatus
    Last Modified in 05/02/19
    286838200004
    [45] † Titanosauridae sp
    Last Modified in 05/02/19
    343252500004
    [46] † Proganochelys quenstedti
    Last Modified in 05/02/19
    141227550017304
    [47] Chelydra serpentina
    Last Modified in 12/05/13
    643041950011004
    [48] Pseudemys scripta
    Last Modified in 12/05/13
    68300208009504
    [49] Geochelone
    Last Modified in 12/05/13
    693011780012404
    [50] Trionyx
    Last Modified in 12/05/13
    7429529400004
    [51] † Baena
    Last Modified in 05/02/19
    2709832007004
    [52] Sphenodon punctatus
    Last Modified in 12/05/13
    553131200019604
    [53] Heloderma
    Last Modified in 12/05/13
    19350339001004
    [54] Corucia zebrata
    Last Modified in 12/05/13
    1535335300004
    [55] Tiliqua
    Last Modified in 12/05/13
    1735335100004
    [56] Cordylus
    Last Modified in 12/05/13
    153551250023204
    [57] Zonosaurus
    Last Modified in 12/05/13
    163551790017804
    [58] Anguis fragilis
    Last Modified in 12/05/13
    812871680012004
    [59] Ophisaurus
    Last Modified in 12/05/13
    7529429300004
    [60] Gerrhonotus
    Last Modified in 12/05/13
    1135835700004
    [61] Elgaria
    Last Modified in 12/05/13
    1535335300004
    [62] † Glyptosaurinae
    Last Modified in 05/02/19
    221147370011304
    [63] † Mesosauridae
    Last Modified in 05/02/19
    19916916900004
    [64] † Millerettidae
    Last Modified in 05/02/19
    16021220800004
    [65] † Anthodon serrarius
    Last Modified in 05/02/19
    12024824800004
    [66] † Scutosaurus karpinskii
    Last Modified in 05/02/19
    9927126900004
    [67] † Bradysaurus seeleyi
    Last Modified in 05/02/19
    10226626600004
    [68] † Pareiasaurus serridens
    Last Modified in 05/02/19
    11025825800004
    [69] † Procolophon
    Last Modified in 05/02/19
    137234840014904
    [70] † Owenetta rubidgei
    Last Modified in 05/02/19
    19817117000004
    [71] † Placodus
    Last Modified in 05/02/19
    1232451350011304
    [72] † Eosauropterygia
    Last Modified in 05/02/19
    22317214500004
    [73] † Cyamodontoidea
    Last Modified in 05/02/19
    165212670013904
    [74] † Rhynchosauria
    Last Modified in 05/02/19
    151226980011904
    [75] † Trilophosaurus buettneri
    Last Modified in 05/02/19
    16820120000004
    [76] † Choristodera
    Last Modified in 05/02/19
    16620320200004
    [77] † Prolacertiformes
    Last Modified in 05/02/19
    15422621400004
    [78] † Acleistorhinus pteroticus
    Last Modified in 05/02/19
    25411411400004
    [79] † Lanthanosuchidae
    Last Modified in 05/02/19
    25112011700004
    [80] † Macroleter poezicus
    Last Modified in 05/02/19
    21815015000004


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M8140 (155 views); M7831 (13 views); M7767 (11 views); M7762 (8 views); M7100 (25 views); M7038 (14 views); M7853 (25 views); M7711 (11 views); M7738 (9 views); M7825 (13 views); M7886 (16 views); M7009 (10 views); M7822 (15 views); M7728 (11 views); M8676 (24 views); M7018 (9 views); M7855 (20 views); M8704 (29 views); M7951 (141 views); M6991 (8 views); M8677 (19 views); M6976 (7 views); M7019 (6 views); M7710 (9 views); M6978 (10 views); M6984 (10 views); M7766 (11 views); M8653 (87 views); M7008 (8 views); M6986 (7 views); M8692 (23 views); M7792 (8 views); M7813 (13 views); M7741 (7 views); M6974 (8 views); M8710 (21 views); M6983 (9 views); M6982 (6 views); M6998 (9 views); M7691 (17 views); M7755 (8 views); M7095 (23 views); M7013 (9 views); M7770 (11 views); M6966 (8 views); M6969 (11 views); M6959 (8 views); M7727 (9 views); M7005 (6 views); M7737 (8 views); M8698 (22 views); M7006 (17 views); M7809 (15 views); M7736 (9 views); M7731 (10 views); M6965 (28 views); M7026 (5 views); M7780 (9 views); M8690 (21 views); M7753 (9 views); M8693 (26 views); M7896 (16 views); M7749 (7 views); M7861 (14 views); M8368 (80 views); M7880 (13 views); M7726 (7 views); M7838 (18 views); M8657 (81 views); M7717 (8 views); M7757 (7 views); M8687 (23 views); M7975 (77 views); M7702 (7 views); M8666 (19 views); M7859 (16 views); M7035 (16 views); M7856 (13 views); M8683 (26 views); M8662 (22 views); M7509 (20 views); M8664 (23 views); M8685 (23 views); M8700 (22 views); M8659 (28 views); M7047 (15 views); M8680 (19 views); M8709 (20 views); M8695 (29 views); M7032 (13 views); M7818 (9 views); M7830 (17 views); M8661 (23 views); M8705 (21 views); M7832 (27 views); M8686 (20 views); M7849 (16 views); M8682 (21 views); M7699 (18 views); M7031 (17 views); M7275 (17 views); M7870 (19 views); M7093 (13 views); M7869 (27 views); M7851 (16 views); M7508 (27 views); M7057 (16 views); M7847 (20 views); M7872 (13 views); M7852 (18 views); M7274 (15 views); M7327 (20 views); M7037 (17 views); M7786 (8 views); M8132 (78 views); M7733 (6 views); M7707 (7 views); M6977 (5 views); M7765 (7 views); M7777 (7 views); M7746 (5 views); M7794 (6 views); M7713 (8 views); M7814 (15 views); M7892 (12 views); M7744 (9 views); M7201 (15 views); M6980 (7 views); M7799 (7 views); M7012 (9 views); M7798 (7 views); M7712 (7 views); M7796 (9 views); M7769 (8 views); M7721 (10 views); M7771 (8 views); M6979 (5 views); M7703 (6 views); M7955 (77 views); M7758 (11 views); M7875 (15 views); M7522 (21 views); M7021 (5 views); M7778 (7 views); M7876 (17 views);
    Matrix views2733Matrix landing page (2290 views); Amniote phylogeny (443 views);
    Project overview5554
    Taxon list20559
    Specimen list191162
    Views for media list9832
    Bibliography455
    Documents list13




    Project downloads

    type number of downloads Individual items downloaded (where applicable)
    Total downloads from project676
    Media downloads132M7063 (1 download); M7067 (3 downloads); M7039 (1 download); M7940 (1 download); M7941 (1 download); M7942 (1 download); M7943 (1 download); M6670 (1 download); M7104 (4 downloads); M6813 (2 downloads); M6951 (1 download); M6779 (3 downloads); M6928 (3 downloads); M6940 (2 downloads); M6953 (1 download); M7059 (1 download); M7081 (1 download); M7082 (1 download); M7899 (3 downloads); M7910 (1 download); M1132 (1 download); M7968 (1 download); M7969 (1 download); M7970 (1 download); M7971 (4 downloads); M7972 (2 downloads); M7973 (2 downloads); M7962 (1 download); M6587 (3 downloads); M7950 (1 download); M7954 (1 download); M7949 (1 download); M249 (2 downloads); M450 (2 downloads); M6983 (1 download); M233 (1 download); M235 (1 download); M241 (1 download); M309 (1 download); M329 (1 download); M330 (1 download); M355 (1 download); M356 (1 download); M358 (1 download); M360 (1 download); M362 (1 download); M365 (1 download); M366 (1 download); M370 (1 download); M371 (1 download); M372 (1 download); M374 (1 download); M376 (1 download); M380 (1 download); M381 (1 download); M382 (1 download); M383 (1 download); M444 (1 download); M445 (1 download); M446 (1 download); M447 (1 download); M448 (1 download); M449 (1 download); M451 (1 download); M452 (1 download); M6588 (1 download); M6673 (1 download); M6674 (1 download); M6859 (1 download); M7886 (2 downloads); M8695 (1 download); M8705 (1 download); M8706 (1 download); M6862 (1 download); M8687 (2 downloads); M8682 (1 download); M7848 (1 download); M7842 (1 download); M7843 (1 download); M7844 (1 download); M7845 (1 download); M7846 (1 download); M7847 (1 download); M7849 (1 download); M7850 (1 download); M7851 (2 downloads); M7852 (1 download); M7853 (1 download); M7854 (1 download); M7855 (1 download); M7874 (1 download); M7875 (1 download); M7876 (1 download); M7877 (1 download); M7878 (1 download); M7879 (1 download); M7880 (1 download); M7881 (1 download); M7882 (1 download); M7883 (1 download); M7884 (1 download); M7885 (1 download); M7887 (1 download); M7889 (1 download); M7890 (1 download); M7891 (1 download); M214 (1 download);
    Matrix downloads81Amniote phylogeny (81 downloads);
    Project downloads463