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: Aetosauridae indet. (MCZ:No #)
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
  • 754 Media
  • 1 Matrix
  • 222 Taxa
  • 428 Specimens
  • 368 Characters
Total size of project's media files: 1042 MB

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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
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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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    Seth Kaufman
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    Robert Hill
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    Mark Miller
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    Maureen O'Leary
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    Taxonomic Overview for Matrix 'M147' (80 Taxa)

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    [1] Seymouriidae
    Last Modified in 12/05/13
    1332351020015004
    [2] Diadectomorpha
    Last Modified in 12/05/13
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    [3] Casea
    Last Modified in 12/05/13
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    [4] Ophiacodon retroversus
    Last Modified in 12/05/13
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    [5] Edaphosauridae
    Last Modified in 12/05/13
    15321621500004
    [6] Sphenacodontidae
    Last Modified in 12/05/13
    13223623600004
    [7] Gorgonopsia
    Last Modified in 12/05/13
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    [8] Procynosuchus
    Last Modified in 12/05/13
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    [9] Morganucodon
    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    [14] Dasypus novemcinctus
    Last Modified in 12/05/13
    78295211008104
    [15] Zaedyus pichiy
    Last Modified in 12/05/13
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    [16] Captorhinidae
    Last Modified in 12/05/13
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    [17] Araeoscelidia
    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    [22] Coahomasuchus kahleorum
    Last Modified in 12/05/13
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    [23] Desmatosuchus haplocerus
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    [24] Typothorax coccinarum
    Last Modified in 12/05/13
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    [25] Paratypothorax andressorum
    Last Modified in 12/05/13
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    [26] Pseudopalatus
    Last Modified in 12/05/13
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    [27] Rutiodon
    Last Modified in 12/05/13
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    [28] Protosuchus richardsoni
    Last Modified in 12/05/13
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    [29] Akanthosuchus langstoni
    Last Modified in 12/05/13
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    [30] Simosuchus clarki
    Last Modified in 12/05/13
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    [31] Goniopholis lucasii
    Last Modified in 12/05/13
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    [32] Mahajangasuchus insignis
    Last Modified in 12/05/13
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    [33] Alligator mississippiensis
    Last Modified in 12/05/13
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    [34] Crocodylus
    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    [36] Gavialis gangeticus
    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    [38] Pinacosaurus grangeri
    Last Modified in 12/05/13
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    [39] Sauropelta edwardsorum
    Last Modified in 12/05/13
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    [40] Stegosaurus stenops
    Last Modified in 12/05/13
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    [41] Scelidosaurus harrisonii
    Last Modified in 12/05/13
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    [42] Scutellosaurus lawleri
    Last Modified in 12/05/13
    25811111000004
    [43] Rapetosaurus krausei
    Last Modified in 12/05/13
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    [44] Saltasaurus loricatus
    Last Modified in 12/05/13
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    [45] Titanosauridae sp
    Last Modified in 12/05/13
    343252500004
    [46] Proganochelys quenstedti
    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    [48] Pseudemys scripta
    Last Modified in 12/05/13
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    [49] Geochelone
    Last Modified in 12/05/13
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    [50] Trionyx
    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    [70] Owenetta rubidgei
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    [71] Placodus
    Last Modified in 12/05/13
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    Last Modified in 12/05/13
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    [73] Cyamodontoidea
    Last Modified in 12/05/13
    165212670013904
    [74] Rhynchosauria
    Last Modified in 12/05/13
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    [75] Trilophosaurus buettneri
    Last Modified in 12/05/13
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    [76] Choristodera
    Last Modified in 12/05/13
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    [77] Prolacertiformes
    Last Modified in 12/05/13
    15422621400004
    [78] Acleistorhinus pteroticus
    Last Modified in 12/05/13
    25411411400004
    [79] Lanthanosuchidae
    Last Modified in 12/05/13
    25112011700004
    [80] Macroleter poezicus
    Last Modified in 12/05/13
    21815015000004


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M7038 (6 views); M7853 (14 views); M7711 (9 views); M7738 (7 views); M7825 (7 views); M7886 (5 views); M7009 (8 views); M7822 (7 views); M7728 (7 views); M8676 (11 views); M7018 (8 views); M7855 (8 views); M8704 (12 views); M7951 (122 views); M6991 (6 views); M8677 (9 views); M6976 (6 views); M7019 (5 views); M7710 (6 views); M6978 (9 views); M6984 (9 views); M7766 (10 views); M8653 (70 views); M7008 (5 views); M6986 (6 views); M8692 (9 views); M7792 (5 views); M7813 (6 views); M7741 (6 views); M6974 (7 views); M8710 (11 views); M6983 (8 views); M6982 (5 views); M6998 (8 views); M7691 (9 views); M7755 (7 views); M7095 (8 views); M7013 (8 views); M7770 (9 views); M6966 (7 views); M6969 (10 views); M6959 (7 views); M7727 (8 views); M7005 (5 views); M7737 (7 views); M8698 (11 views); M7006 (16 views); M7809 (8 views); M7736 (8 views); M7731 (6 views); M6965 (25 views); M7026 (5 views); M7780 (7 views); M8690 (7 views); M7753 (8 views); M8693 (15 views); M7896 (7 views); M7749 (6 views); M7861 (6 views); M8368 (67 views); M7880 (6 views); M7726 (5 views); M7838 (9 views); M8657 (66 views); M7717 (6 views); M7757 (6 views); M8687 (8 views); M7975 (64 views); M7702 (6 views); M8666 (7 views); M7859 (9 views); M7035 (7 views); M7856 (6 views); M8683 (10 views); M8662 (9 views); M7509 (6 views); M8664 (10 views); M8685 (9 views); M8700 (7 views); M8659 (7 views); M7047 (5 views); M8680 (7 views); M8709 (7 views); M8695 (10 views); M7032 (5 views); M7818 (5 views); M7830 (9 views); M8661 (8 views); M8705 (8 views); M7832 (19 views); M8686 (7 views); M7849 (8 views); M8682 (7 views); M7699 (7 views); M7031 (6 views); M7275 (8 views); M7870 (9 views); M7093 (4 views); M7869 (8 views); M7851 (7 views); M7508 (10 views); M7057 (6 views); M7847 (9 views); M7872 (5 views); M7852 (6 views); M7274 (7 views); M7327 (7 views); M7037 (7 views); M7786 (4 views); M8132 (65 views); M7733 (5 views); M7707 (6 views); M6977 (4 views); M7765 (6 views); M7777 (5 views); M7746 (4 views); M7794 (4 views); M7713 (6 views); M7814 (6 views); M7892 (5 views); M7744 (7 views); M7201 (6 views); M6980 (6 views); M7799 (4 views); M7012 (7 views); M7798 (5 views); M7712 (6 views); M7796 (7 views); M7769 (5 views); M7721 (8 views); M7771 (5 views); M6979 (4 views); M7703 (5 views); M7955 (61 views); M7758 (8 views); M7875 (7 views); M7522 (7 views); M7021 (4 views); M7778 (6 views); M7876 (5 views);
    Matrix views1688Matrix landing page (1386 views); Amniote phylogeny (302 views);
    Project overview3430
    Taxon list10321
    Specimen list109632
    Views for media list5593




    Project downloads

    type number of downloads Individual items downloaded (where applicable)
    Total downloads from project401
    Media downloads90M7063 (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);
    Matrix downloads65Amniote phylogeny (65 downloads);
    Project downloads246