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 - Media downloads: 141
- Matrix downloads: 97
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 Taxon name last Modified on 05/02/19 | 133 | 235 | 102 | 0 | 0 | 150 | 0 | 4 |
[2] † Diadectomorpha Taxon name last Modified on 05/02/19 | 151 | 224 | 77 | 0 | 0 | 143 | 0 | 4 |
[3] † Casea Taxon name last Modified on 05/02/19 | 142 | 226 | 226 | 0 | 0 | 0 | 0 | 4 |
[4] † Ophiacodon retroversus Taxon name last Modified on 05/02/19 | 137 | 231 | 85 | 0 | 0 | 146 | 0 | 4 |
[5] † Edaphosauridae Taxon name last Modified on 05/02/19 | 153 | 216 | 215 | 0 | 0 | 0 | 0 | 4 |
[6] † Sphenacodontidae Taxon name last Modified on 05/02/19 | 132 | 236 | 236 | 0 | 0 | 0 | 0 | 4 |
[7] † Gorgonopsia Taxon name last Modified on 05/02/19 | 145 | 223 | 223 | 0 | 0 | 0 | 0 | 4 |
[8] † Procynosuchus Taxon name last Modified on 05/02/19 | 134 | 236 | 234 | 0 | 0 | 0 | 0 | 4 |
[9] † Morganucodon Taxon name last Modified on 05/02/19 | 175 | 193 | 193 | 0 | 0 | 0 | 0 | 4 |
[10] † Glyptodon reticulatus Taxon name last Modified on 05/02/19 | 143 | 226 | 208 | 0 | 0 | 22 | 0 | 4 |
[11] † Glyptotherium Taxon name last Modified on 05/02/19 | 148 | 222 | 220 | 0 | 0 | 0 | 0 | 4 |
[12] † Doedicurus giganteus Taxon name last Modified on 05/02/19 | 144 | 224 | 207 | 0 | 0 | 17 | 0 | 4 |
[13] † Panochthus frenzelianus Taxon name last Modified on 05/02/19 | 143 | 226 | 208 | 0 | 0 | 17 | 0 | 4 |
[14] Dasypus novemcinctus Taxon name last Modified on 12/05/13 | 78 | 295 | 211 | 0 | 0 | 81 | 0 | 4 |
[15] Zaedyus pichiy Taxon name last Modified on 12/05/13 | 78 | 295 | 290 | 0 | 0 | 0 | 0 | 4 |
[16] † Captorhinidae Taxon name last Modified on 05/02/19 | 131 | 239 | 118 | 0 | 0 | 121 | 0 | 4 |
[17] † Araeoscelidia Taxon name last Modified on 05/02/19 | 134 | 239 | 7 | 0 | 0 | 232 | 0 | 4 |
[18] † Paleothyris acadiana Taxon name last Modified on 05/02/19 | 153 | 215 | 215 | 0 | 0 | 0 | 0 | 4 |
[19] † Claudiosaurus germaini Taxon name last Modified on 05/02/19 | 184 | 184 | 184 | 0 | 0 | 0 | 0 | 4 |
[20] † Younginiformes Taxon name last Modified on 05/02/19 | 165 | 210 | 203 | 0 | 0 | 0 | 0 | 4 |
[21] † Kuehneosauridae Taxon name last Modified on 05/02/19 | 248 | 120 | 120 | 0 | 0 | 0 | 0 | 4 |
[22] † Coahomasuchus kahleorum Taxon name last Modified on 05/02/19 | 282 | 86 | 86 | 0 | 0 | 0 | 0 | 4 |
[23] † Desmatosuchus haplocerus Taxon name last Modified on 05/02/19 | 148 | 221 | 153 | 0 | 0 | 78 | 0 | 4 |
[24] † Typothorax coccinarum Taxon name last Modified on 05/02/19 | 228 | 140 | 119 | 0 | 0 | 25 | 0 | 4 |
[25] † Paratypothorax andressorum Taxon name last Modified on 05/02/19 | 317 | 51 | 51 | 0 | 0 | 0 | 0 | 4 |
[26] † Pseudopalatus Taxon name last Modified on 05/02/19 | 156 | 214 | 85 | 0 | 0 | 136 | 0 | 4 |
[27] † Rutiodon Taxon name last Modified on 05/02/19 | 128 | 243 | 191 | 0 | 0 | 53 | 0 | 4 |
[28] † Protosuchus richardsoni Taxon name last Modified on 05/02/19 | 142 | 226 | 169 | 0 | 0 | 58 | 0 | 4 |
[29] † Akanthosuchus langstoni Taxon name last Modified on 05/02/19 | 313 | 58 | 55 | 0 | 0 | 0 | 0 | 4 |
[30] † Simosuchus clarki Taxon name last Modified on 05/02/19 | 172 | 197 | 47 | 0 | 0 | 151 | 0 | 4 |
[31] † Goniopholis lucasii Taxon name last Modified on 05/02/19 | 205 | 163 | 49 | 0 | 0 | 114 | 0 | 4 |
[32] † Mahajangasuchus insignis Taxon name last Modified on 05/02/19 | 164 | 204 | 26 | 0 | 0 | 182 | 0 | 4 |
[33] Alligator mississippiensis Taxon name last Modified on 12/05/13 | 15 | 354 | 135 | 0 | 0 | 229 | 0 | 4 |
[34] Crocodylus Taxon name last Modified on 12/05/13 | 17 | 352 | 194 | 0 | 0 | 161 | 0 | 4 |
[35] Caiman crocodilus Taxon name last Modified on 12/05/13 | 18 | 350 | 227 | 0 | 0 | 123 | 0 | 4 |
[36] Gavialis gangeticus Taxon name last Modified on 12/05/13 | 18 | 350 | 225 | 0 | 0 | 127 | 0 | 4 |
[37] † Euoplocephalus tutus Taxon name last Modified on 05/02/19 | 155 | 214 | 213 | 0 | 0 | 0 | 0 | 4 |
[38] † Pinacosaurus grangeri Taxon name last Modified on 05/02/19 | 166 | 202 | 76 | 0 | 0 | 129 | 0 | 4 |
[39] † Sauropelta edwardsorum Taxon name last Modified on 05/02/19 | 154 | 214 | 214 | 0 | 0 | 0 | 0 | 4 |
[40] † Stegosaurus stenops Taxon name last Modified on 05/02/19 | 129 | 239 | 239 | 0 | 0 | 0 | 0 | 4 |
[41] † Scelidosaurus harrisonii Taxon name last Modified on 05/02/19 | 201 | 168 | 167 | 0 | 0 | 0 | 0 | 4 |
[42] † Scutellosaurus lawleri Taxon name last Modified on 05/02/19 | 258 | 111 | 110 | 0 | 0 | 0 | 0 | 4 |
[43] † Rapetosaurus krausei Taxon name last Modified on 05/02/19 | 342 | 26 | 8 | 0 | 0 | 18 | 0 | 4 |
[44] † Saltasaurus loricatus Taxon name last Modified on 05/02/19 | 286 | 83 | 82 | 0 | 0 | 0 | 0 | 4 |
[45] † Titanosauridae sp Taxon name last Modified on 05/02/19 | 343 | 25 | 25 | 0 | 0 | 0 | 0 | 4 |
[46] † Proganochelys quenstedti Taxon name last Modified on 05/02/19 | 141 | 227 | 55 | 0 | 0 | 173 | 0 | 4 |
[47] Chelydra serpentina Taxon name last Modified on 12/05/13 | 64 | 304 | 195 | 0 | 0 | 110 | 0 | 4 |
[48] Pseudemys scripta Taxon name last Modified on 12/05/13 | 68 | 300 | 208 | 0 | 0 | 95 | 0 | 4 |
[49] Geochelone Taxon name last Modified on 12/05/13 | 69 | 301 | 178 | 0 | 0 | 124 | 0 | 4 |
[50] Trionyx Taxon name last Modified on 12/05/13 | 74 | 295 | 294 | 0 | 0 | 0 | 0 | 4 |
[51] † Baena Taxon name last Modified on 05/02/19 | 270 | 98 | 32 | 0 | 0 | 70 | 0 | 4 |
[52] Sphenodon punctatus Taxon name last Modified on 12/05/13 | 55 | 313 | 120 | 0 | 0 | 196 | 0 | 4 |
[53] Heloderma Taxon name last Modified on 12/05/13 | 19 | 350 | 339 | 0 | 0 | 10 | 0 | 4 |
[54] Corucia zebrata Taxon name last Modified on 12/05/13 | 15 | 353 | 353 | 0 | 0 | 0 | 0 | 4 |
[55] Tiliqua Taxon name last Modified on 12/05/13 | 17 | 353 | 351 | 0 | 0 | 0 | 0 | 4 |
[56] Cordylus Taxon name last Modified on 12/05/13 | 15 | 355 | 125 | 0 | 0 | 232 | 0 | 4 |
[57] Zonosaurus Taxon name last Modified on 12/05/13 | 16 | 355 | 179 | 0 | 0 | 178 | 0 | 4 |
[58] Anguis fragilis Taxon name last Modified on 12/05/13 | 81 | 287 | 168 | 0 | 0 | 120 | 0 | 4 |
[59] Ophisaurus Taxon name last Modified on 12/05/13 | 75 | 294 | 293 | 0 | 0 | 0 | 0 | 4 |
[60] Gerrhonotus Taxon name last Modified on 12/05/13 | 11 | 358 | 357 | 0 | 0 | 0 | 0 | 4 |
[61] Elgaria Taxon name last Modified on 12/05/13 | 15 | 353 | 353 | 0 | 0 | 0 | 0 | 4 |
[62] † Glyptosaurinae Taxon name last Modified on 05/02/19 | 221 | 147 | 37 | 0 | 0 | 113 | 0 | 4 |
[63] † Mesosauridae Taxon name last Modified on 05/02/19 | 199 | 169 | 169 | 0 | 0 | 0 | 0 | 4 |
[64] † Millerettidae Taxon name last Modified on 05/02/19 | 160 | 212 | 208 | 0 | 0 | 0 | 0 | 4 |
[65] † Anthodon serrarius Taxon name last Modified on 05/02/19 | 120 | 248 | 248 | 0 | 0 | 0 | 0 | 4 |
[66] † Scutosaurus karpinskii Taxon name last Modified on 05/02/19 | 99 | 271 | 269 | 0 | 0 | 0 | 0 | 4 |
[67] † Bradysaurus seeleyi Taxon name last Modified on 05/02/19 | 102 | 266 | 266 | 0 | 0 | 0 | 0 | 4 |
[68] † Pareiasaurus serridens Taxon name last Modified on 05/02/19 | 110 | 258 | 258 | 0 | 0 | 0 | 0 | 4 |
[69] † Procolophon Taxon name last Modified on 05/02/19 | 137 | 234 | 84 | 0 | 0 | 149 | 0 | 4 |
[70] † Owenetta rubidgei Taxon name last Modified on 05/02/19 | 198 | 171 | 170 | 0 | 0 | 0 | 0 | 4 |
[71] † Placodus Taxon name last Modified on 05/02/19 | 123 | 245 | 135 | 0 | 0 | 113 | 0 | 4 |
[72] † Eosauropterygia Taxon name last Modified on 05/02/19 | 223 | 172 | 145 | 0 | 0 | 0 | 0 | 4 |
[73] † Cyamodontoidea Taxon name last Modified on 05/02/19 | 165 | 212 | 67 | 0 | 0 | 139 | 0 | 4 |
[74] † Rhynchosauria Taxon name last Modified on 05/02/19 | 151 | 226 | 98 | 0 | 0 | 119 | 0 | 4 |
[75] † Trilophosaurus buettneri Taxon name last Modified on 05/02/19 | 168 | 201 | 200 | 0 | 0 | 0 | 0 | 4 |
[76] † Choristodera Taxon name last Modified on 05/02/19 | 166 | 203 | 202 | 0 | 0 | 0 | 0 | 4 |
[77] † Prolacertiformes Taxon name last Modified on 05/02/19 | 154 | 226 | 214 | 0 | 0 | 0 | 0 | 4 |
[78] † Acleistorhinus pteroticus Taxon name last Modified on 05/02/19 | 254 | 114 | 114 | 0 | 0 | 0 | 0 | 4 |
[79] † Lanthanosuchidae Taxon name last Modified on 05/02/19 | 251 | 120 | 117 | 0 | 0 | 0 | 0 | 4 |
[80] † Macroleter poezicus Taxon name last Modified on 05/02/19 | 218 | 150 | 150 | 0 | 0 | 0 | 0 | 4 |
Project downloads
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 699 | |
Media downloads | 141 | 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 (2 downloads); 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 (2 downloads); 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); M7967 (1 download); M7966 (1 download); M7925 (1 download); M6955 (1 download); M6957 (1 download); M6956 (1 download); M6958 (1 download); |
Matrix downloads | 97 | Amniote phylogeny (97 downloads); |
Project downloads | 461 |