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.
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.]Read the article »
Project DOI: 10.7934/P46, http://dx.doi.org/10.7934/P46
This project contains | Matrices |
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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
MorphoBank Project 46
- Creation Date:
01 June 2006 - Publication Date:
01 January 2007 - Project views: 274256
- Media downloads: 99
- Matrix downloads: 72
This research
supported by
Authors' Institutions
- Stony Brook University
Members
member name | taxa ![]() |
specimens ![]() |
media ![]() | media notes | chars ![]() | character
| cell scorings (scored, NPA, "-") | cell
| rules ![]() | ||||||||||
Seth Kaufman Project Administrator | 0 | 0 | 0 | 0 | 368 | 0 | 0 | 0 | 17947 (17947, 0, 0) | 0 | 0 | 0 | 0 | ||||||
MorphoBank Curator | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | 0 | ||||||
Robert Hill Full membership ![]() | 139 | 37 | 673 | 10 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | 0 | ||||||
Mark Miller Full membership ![]() | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | 0 | ||||||
Maureen O'Leary Full membership ![]() | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M147' (80 Taxa)
taxon | unscored cells |
scored cells ![]() |
no cell support ![]() |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] † Seymouriidae Last Modified in 05/02/19 | 133 | 235 | 102 | 0 | 0 | 150 | 0 | 4 ![]() |
[2] † Diadectomorpha Last Modified in 05/02/19 | 151 | 224 | 77 | 0 | 0 | 143 | 0 | 4 ![]() |
[3] † Casea Last Modified in 05/02/19 | 142 | 226 | 226 | 0 | 0 | 0 | 0 | 4 ![]() |
[4] † Ophiacodon retroversus Last Modified in 05/02/19 | 137 | 231 | 85 | 0 | 0 | 146 | 0 | 4 ![]() |
[5] † Edaphosauridae Last Modified in 05/02/19 | 153 | 216 | 215 | 0 | 0 | 0 | 0 | 4 ![]() |
[6] † Sphenacodontidae Last Modified in 05/02/19 | 132 | 236 | 236 | 0 | 0 | 0 | 0 | 4 ![]() |
[7] † Gorgonopsia Last Modified in 05/02/19 | 145 | 223 | 223 | 0 | 0 | 0 | 0 | 4 ![]() |
[8] † Procynosuchus Last Modified in 05/02/19 | 134 | 236 | 234 | 0 | 0 | 0 | 0 | 4 ![]() |
[9] † Morganucodon Last Modified in 05/02/19 | 175 | 193 | 193 | 0 | 0 | 0 | 0 | 4 ![]() |
[10] † Glyptodon reticulatus Last Modified in 05/02/19 | 143 | 226 | 208 | 0 | 0 | 22 | 0 | 4 ![]() |
[11] † Glyptotherium Last Modified in 05/02/19 | 148 | 222 | 220 | 0 | 0 | 0 | 0 | 4 ![]() |
[12] † Doedicurus giganteus Last Modified in 05/02/19 | 144 | 224 | 207 | 0 | 0 | 17 | 0 | 4 ![]() |
[13] † Panochthus frenzelianus Last Modified in 05/02/19 | 143 | 226 | 208 | 0 | 0 | 17 | 0 | 4 ![]() |
[14] Dasypus novemcinctus Last Modified in 12/05/13 | 78 | 295 | 211 | 0 | 0 | 81 | 0 | 4 ![]() |
[15] Zaedyus pichiy Last Modified in 12/05/13 | 78 | 295 | 290 | 0 | 0 | 0 | 0 | 4 ![]() |
[16] † Captorhinidae Last Modified in 05/02/19 | 131 | 239 | 118 | 0 | 0 | 121 | 0 | 4 ![]() |
[17] † Araeoscelidia Last Modified in 05/02/19 | 134 | 239 | 7 | 0 | 0 | 232 | 0 | 4 ![]() |
[18] † Paleothyris acadiana Last Modified in 05/02/19 | 153 | 215 | 215 | 0 | 0 | 0 | 0 | 4 ![]() |
[19] † Claudiosaurus germaini Last Modified in 05/02/19 | 184 | 184 | 184 | 0 | 0 | 0 | 0 | 4 ![]() |
[20] † Younginiformes Last Modified in 05/02/19 | 165 | 210 | 203 | 0 | 0 | 0 | 0 | 4 ![]() |
[21] † Kuehneosauridae Last Modified in 05/02/19 | 248 | 120 | 120 | 0 | 0 | 0 | 0 | 4 ![]() |
[22] † Coahomasuchus kahleorum Last Modified in 05/02/19 | 282 | 86 | 86 | 0 | 0 | 0 | 0 | 4 ![]() |
[23] † Desmatosuchus haplocerus Last Modified in 05/02/19 | 148 | 221 | 153 | 0 | 0 | 78 | 0 | 4 ![]() |
[24] † Typothorax coccinarum Last Modified in 05/02/19 | 228 | 140 | 119 | 0 | 0 | 25 | 0 | 4 ![]() |
[25] † Paratypothorax andressorum Last Modified in 05/02/19 | 317 | 51 | 51 | 0 | 0 | 0 | 0 | 4 ![]() |
[26] † Pseudopalatus Last Modified in 05/02/19 | 156 | 214 | 85 | 0 | 0 | 136 | 0 | 4 ![]() |
[27] † Rutiodon Last Modified in 05/02/19 | 128 | 243 | 191 | 0 | 0 | 53 | 0 | 4 ![]() |
[28] † Protosuchus richardsoni Last Modified in 05/02/19 | 142 | 226 | 169 | 0 | 0 | 58 | 0 | 4 ![]() |
[29] † Akanthosuchus langstoni Last Modified in 05/02/19 | 313 | 58 | 55 | 0 | 0 | 0 | 0 | 4 ![]() |
[30] † Simosuchus clarki Last Modified in 05/02/19 | 172 | 197 | 47 | 0 | 0 | 151 | 0 | 4 ![]() |
[31] † Goniopholis lucasii Last Modified in 05/02/19 | 205 | 163 | 49 | 0 | 0 | 114 | 0 | 4 ![]() |
[32] † Mahajangasuchus insignis Last Modified in 05/02/19 | 164 | 204 | 26 | 0 | 0 | 182 | 0 | 4 ![]() |
[33] Alligator mississippiensis Last Modified in 12/05/13 | 15 | 354 | 135 | 0 | 0 | 229 | 0 | 4 ![]() |
[34] Crocodylus Last Modified in 12/05/13 | 17 | 352 | 194 | 0 | 0 | 161 | 0 | 4 ![]() |
[35] Caiman crocodilus Last Modified in 12/05/13 | 18 | 350 | 227 | 0 | 0 | 123 | 0 | 4 ![]() |
[36] Gavialis gangeticus Last Modified in 12/05/13 | 18 | 350 | 225 | 0 | 0 | 127 | 0 | 4 ![]() |
[37] † Euoplocephalus tutus Last Modified in 05/02/19 | 155 | 214 | 213 | 0 | 0 | 0 | 0 | 4 ![]() |
[38] † Pinacosaurus grangeri Last Modified in 05/02/19 | 166 | 202 | 76 | 0 | 0 | 129 | 0 | 4 ![]() |
[39] † Sauropelta edwardsorum Last Modified in 05/02/19 | 154 | 214 | 214 | 0 | 0 | 0 | 0 | 4 ![]() |
[40] † Stegosaurus stenops Last Modified in 05/02/19 | 129 | 239 | 239 | 0 | 0 | 0 | 0 | 4 ![]() |
[41] † Scelidosaurus harrisonii Last Modified in 05/02/19 | 201 | 168 | 167 | 0 | 0 | 0 | 0 | 4 ![]() |
[42] † Scutellosaurus lawleri Last Modified in 05/02/19 | 258 | 111 | 110 | 0 | 0 | 0 | 0 | 4 ![]() |
[43] † Rapetosaurus krausei Last Modified in 05/02/19 | 342 | 26 | 8 | 0 | 0 | 18 | 0 | 4 ![]() |
[44] † Saltasaurus loricatus Last Modified in 05/02/19 | 286 | 83 | 82 | 0 | 0 | 0 | 0 | 4 ![]() |
[45] † Titanosauridae sp Last Modified in 05/02/19 | 343 | 25 | 25 | 0 | 0 | 0 | 0 | 4 ![]() |
[46] † Proganochelys quenstedti Last Modified in 05/02/19 | 141 | 227 | 55 | 0 | 0 | 173 | 0 | 4 ![]() |
[47] Chelydra serpentina Last Modified in 12/05/13 | 64 | 304 | 195 | 0 | 0 | 110 | 0 | 4 ![]() |
[48] Pseudemys scripta Last Modified in 12/05/13 | 68 | 300 | 208 | 0 | 0 | 95 | 0 | 4 ![]() |
[49] Geochelone Last Modified in 12/05/13 | 69 | 301 | 178 | 0 | 0 | 124 | 0 | 4 ![]() |
[50] Trionyx Last Modified in 12/05/13 | 74 | 295 | 294 | 0 | 0 | 0 | 0 | 4 ![]() |
[51] † Baena Last Modified in 05/02/19 | 270 | 98 | 32 | 0 | 0 | 70 | 0 | 4 ![]() |
[52] Sphenodon punctatus Last Modified in 12/05/13 | 55 | 313 | 120 | 0 | 0 | 196 | 0 | 4 ![]() |
[53] Heloderma Last Modified in 12/05/13 | 19 | 350 | 339 | 0 | 0 | 10 | 0 | 4 ![]() |
[54] Corucia zebrata Last Modified in 12/05/13 | 15 | 353 | 353 | 0 | 0 | 0 | 0 | 4 ![]() |
[55] Tiliqua Last Modified in 12/05/13 | 17 | 353 | 351 | 0 | 0 | 0 | 0 | 4 ![]() |
[56] Cordylus Last Modified in 12/05/13 | 15 | 355 | 125 | 0 | 0 | 232 | 0 | 4 ![]() |
[57] Zonosaurus Last Modified in 12/05/13 | 16 | 355 | 179 | 0 | 0 | 178 | 0 | 4 ![]() |
[58] Anguis fragilis Last Modified in 12/05/13 | 81 | 287 | 168 | 0 | 0 | 120 | 0 | 4 ![]() |
[59] Ophisaurus Last Modified in 12/05/13 | 75 | 294 | 293 | 0 | 0 | 0 | 0 | 4 ![]() |
[60] Gerrhonotus Last Modified in 12/05/13 | 11 | 358 | 357 | 0 | 0 | 0 | 0 | 4 ![]() |
[61] Elgaria Last Modified in 12/05/13 | 15 | 353 | 353 | 0 | 0 | 0 | 0 | 4 ![]() |
[62] † Glyptosaurinae Last Modified in 05/02/19 | 221 | 147 | 37 | 0 | 0 | 113 | 0 | 4 ![]() |
[63] † Mesosauridae Last Modified in 05/02/19 | 199 | 169 | 169 | 0 | 0 | 0 | 0 | 4 ![]() |
[64] † Millerettidae Last Modified in 05/02/19 | 160 | 212 | 208 | 0 | 0 | 0 | 0 | 4 ![]() |
[65] † Anthodon serrarius Last Modified in 05/02/19 | 120 | 248 | 248 | 0 | 0 | 0 | 0 | 4 ![]() |
[66] † Scutosaurus karpinskii Last Modified in 05/02/19 | 99 | 271 | 269 | 0 | 0 | 0 | 0 | 4 ![]() |
[67] † Bradysaurus seeleyi Last Modified in 05/02/19 | 102 | 266 | 266 | 0 | 0 | 0 | 0 | 4 ![]() |
[68] † Pareiasaurus serridens Last Modified in 05/02/19 | 110 | 258 | 258 | 0 | 0 | 0 | 0 | 4 ![]() |
[69] † Procolophon Last Modified in 05/02/19 | 137 | 234 | 84 | 0 | 0 | 149 | 0 | 4 ![]() |
[70] † Owenetta rubidgei Last Modified in 05/02/19 | 198 | 171 | 170 | 0 | 0 | 0 | 0 | 4 ![]() |
[71] † Placodus Last Modified in 05/02/19 | 123 | 245 | 135 | 0 | 0 | 113 | 0 | 4 ![]() |
[72] † Eosauropterygia Last Modified in 05/02/19 | 223 | 172 | 145 | 0 | 0 | 0 | 0 | 4 ![]() |
[73] † Cyamodontoidea Last Modified in 05/02/19 | 165 | 212 | 67 | 0 | 0 | 139 | 0 | 4 ![]() |
[74] † Rhynchosauria Last Modified in 05/02/19 | 151 | 226 | 98 | 0 | 0 | 119 | 0 | 4 ![]() |
[75] † Trilophosaurus buettneri Last Modified in 05/02/19 | 168 | 201 | 200 | 0 | 0 | 0 | 0 | 4 ![]() |
[76] † Choristodera Last Modified in 05/02/19 | 166 | 203 | 202 | 0 | 0 | 0 | 0 | 4 ![]() |
[77] † Prolacertiformes Last Modified in 05/02/19 | 154 | 226 | 214 | 0 | 0 | 0 | 0 | 4 ![]() |
[78] † Acleistorhinus pteroticus Last Modified in 05/02/19 | 254 | 114 | 114 | 0 | 0 | 0 | 0 | 4 ![]() |
[79] † Lanthanosuchidae Last Modified in 05/02/19 | 251 | 120 | 117 | 0 | 0 | 0 | 0 | 4 ![]() |
[80] † Macroleter poezicus Last Modified in 05/02/19 | 218 | 150 | 150 | 0 | 0 | 0 | 0 | 4 ![]() |
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M7732 (7 views); M8689 (16 views); M6958 (5 views); M7774 (8 views); M7002 (6 views); M7759 (7 views); M7874 (14 views); M7877 (9 views); M8694 (16 views); M7684 (14 views); M7698 (15 views); M7272 (24 views); M7017 (8 views); M7857 (11 views); M7034 (18 views); M7773 (6 views); M7027 (8 views); M7841 (9 views); M7823 (11 views); M7816 (6 views); M7819 (9 views); M6987 (8 views); M7788 (8 views); M7085 (13 views); M7833 (8 views); M7797 (9 views); M7854 (11 views); M7781 (8 views); M7101 (15 views); M7044 (10 views); M8708 (10 views); M7743 (7 views); M7820 (9 views); M7751 (8 views); M8703 (16 views); M7328 (15 views); M8126 (64 views); M6981 (9 views); M7049 (11 views); M7961 (60 views); M7004 (5 views); M7016 (6 views); M8699 (12 views); M6960 (5 views); M7834 (9 views); M7734 (6 views); M7815 (9 views); M7843 (9 views); M7686 (13 views); M7760 (8 views); M7719 (6 views); M7742 (6 views); M7824 (9 views); M7690 (13 views); M7964 (57 views); M7103 (12 views); M8127 (75 views); M7730 (8 views); M8668 (19 views); M7966 (111 views); M8678 (12 views); M7827 (9 views); M8375 (59 views); M7772 (6 views); M7935 (73 views); M7029 (9 views); M7723 (5 views); M7033 (13 views); M8369 (84 views); M7957 (83 views); M8371 (153 views); M7976 (131 views); M8378 (95 views); M7947 (66 views); M7924 (131 views); M8656 (65 views); M8376 (72 views); M7030 (13 views); M7269 (12 views); M7936 (53 views); M7519 (12 views); M7040 (9 views); M7931 (113 views); M7705 (9 views); M7326 (11 views); M7916 (105 views); M7685 (14 views); M7088 (15 views); M7036 (12 views); M8696 (11 views); M7918 (127 views); M7810 (11 views); M7858 (8 views); M7846 (7 views); M7837 (10 views); M7842 (22 views); M7092 (15 views); M8660 (12 views); M7020 (7 views); M7867 (9 views); M7693 (11 views); M7885 (10 views); M7812 (11 views); M7888 (13 views); M7883 (12 views); M7277 (13 views); M7001 (8 views); M8681 (11 views); M8665 (15 views); M7048 (9 views); M8684 (15 views); M8140 (149 views); M7831 (9 views); M7767 (10 views); M7762 (7 views); M7100 (16 views); M7038 (8 views); M7853 (15 views); M7711 (9 views); M7738 (7 views); M7825 (8 views); M7886 (8 views); M7009 (9 views); M7822 (9 views); M7728 (7 views); M8676 (14 views); M7018 (8 views); M7855 (9 views); M8704 (16 views); M7951 (125 views); M6991 (7 views); M8677 (12 views); M6976 (6 views); M7019 (5 views); M7710 (6 views); M6978 (9 views); M6984 (9 views); M7766 (10 views); M8653 (74 views); M7008 (6 views); M6986 (6 views); M8692 (15 views); M7792 (5 views); M7813 (9 views); M7741 (6 views); M6974 (7 views); M8710 (13 views); M6983 (8 views); M6982 (5 views); M6998 (8 views); M7691 (13 views); M7755 (7 views); M7095 (14 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 (13 views); M7006 (16 views); M7809 (10 views); M7736 (8 views); M7731 (6 views); M6965 (27 views); M7026 (5 views); M7780 (7 views); M8690 (11 views); M7753 (8 views); M8693 (17 views); M7896 (9 views); M7749 (6 views); M7861 (7 views); M8368 (70 views); M7880 (7 views); M7726 (5 views); M7838 (11 views); M8657 (70 views); M7717 (6 views); M7757 (6 views); M8687 (13 views); M7975 (69 views); M7702 (6 views); M8666 (9 views); M7859 (10 views); M7035 (12 views); M7856 (7 views); M8683 (14 views); M8662 (13 views); M7509 (9 views); M8664 (14 views); M8685 (14 views); M8700 (10 views); M8659 (10 views); M7047 (8 views); M8680 (11 views); M8709 (10 views); M8695 (14 views); M7032 (9 views); M7818 (6 views); M7830 (10 views); M8661 (10 views); M8705 (12 views); M7832 (21 views); M8686 (11 views); M7849 (9 views); M8682 (10 views); M7699 (13 views); M7031 (11 views); M7275 (12 views); M7870 (11 views); M7093 (6 views); M7869 (9 views); M7851 (8 views); M7508 (22 views); M7057 (8 views); M7847 (10 views); M7872 (6 views); M7852 (7 views); M7274 (10 views); M7327 (11 views); M7037 (13 views); M7786 (4 views); M8132 (69 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 (8 views); M7892 (6 views); M7744 (7 views); M7201 (9 views); M6980 (6 views); M7799 (5 views); M7012 (8 views); M7798 (6 views); M7712 (6 views); M7796 (8 views); M7769 (5 views); M7721 (8 views); M7771 (5 views); M6979 (4 views); M7703 (5 views); M7955 (64 views); M7758 (8 views); M7875 (8 views); M7522 (10 views); M7021 (4 views); M7778 (6 views); M7876 (7 views); |
Matrix views | 2295 | Matrix landing page (1925 views); Amniote phylogeny (370 views); |
Project overview | 4582 | |
Taxon list | 16539 | |
Specimen list | 142402 | |
Views for media list | 7499 | |
Bibliography | 210 | |
Documents list | 1 |
Project downloads 
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 517 | |
Media downloads | 99 | M7063 (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 (1 download); M8695 (1 download); M8705 (1 download); M8706 (1 download); M6862 (1 download); M8687 (2 downloads); M8682 (1 download); |
Matrix downloads | 72 | Amniote phylogeny (72 downloads); |
Project downloads | 346 |