Project 790: R. Bieler, P. N. Mikkelsen, T. M. Collins, E. A. Glover, V. L. González, D. L. Graf, E. M. Harper, J. Healy, G. Y. Kawauchi, P. P. Sharma, S. Staubach, E. E. Strong, J. D. Taylor, I. Tëmkin, J. D. Zardus, S. Clark, A. Guzmán, E. McIntyre, P. Sharp, G. Giribet. 2014. Investigating the Bivalve Tree of Life—an exemplar-based approach combining molecular and novel morphological characters. Invertebrate Systematics. 28 (1):32-115.
Abstract
To re-evaluate the relationships of the major bivalve lineages, we amassed detailed morpho-anatomical, ultrastructural and molecular sequence data for a targeted selection of exemplar bivalves spanning the phylogenetic diversity of the class. We included molecular data for 103 bivalve species (up to five markers) and also analysed a subset of taxa with four additional nuclear protein-encoding genes. Novel as well as historically employed morphological characters were explored, and we systematically disassembled widely used descriptors such as gill and stomach ‘types’. Phylogenetic analyses, conducted using parsimony direct optimisation and probabilistic methods on static alignments (maximum likelihood and Bayesian inference) of the molecular data, both alone and in combination with morphological characters, offer a robust test of bivalve relationships. A calibrated phylogeny also provided insights into the tempo of bivalve evolution. Finally, an analysis of the informativeness of morphological characters showed that sperm ultrastructure characters are among the best morphological features to diagnose bivalve clades, followed by characters of the shell, including its microstructure. Our study found support for monophyly of most broadly recognised higher bivalve taxa, although support was not uniform for Protobranchia. However, monophyly of the bivalves with protobranchiate gills was the best-supported hypothesis with incremental morphological and/or molecular sequence data. Autobranchia, Pteriomorphia, Heteroconchia, Palaeoheterodonta, Archiheterodonta, Euheterodonta, Anomalodesmata and Imparidentia new clade ( = Euheterodonta excluding Anomalodesmata) were recovered across analyses, irrespective of data treatment or analytical framework. Another clade supported by our analyses but not formally recognised in the literature includes Palaeoheterodonta and Archiheterodonta, which emerged under multiple analytical conditions. The origin and diversification of each of these major clades is Cambrian or Ordovician, except for Archiheterodonta, which diverged from Palaeoheterodonta during the Cambrian, but diversified during the Mesozoic. Although the radiation of some lineages was shifted towards the Palaeozoic (Pteriomorphia, Anomalodesmata), or presented a gap between origin and diversification (Archiheterodonta, Unionida), Imparidentia showed steady diversification through the Palaeozoic and Mesozoic. Finally, a classification system with six major monophyletic lineages is proposed to comprise modern Bivalvia: Protobranchia, Pteriomorphia, Palaeoheterodonta, Archiheterodonta, Anomalodesmata and Imparidentia.Read the article »
Project DOI: 10.7934/P790, http://dx.doi.org/10.7934/P790
This project contains | Matrices |
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Download Project SDD File | Total scored cells: 19904 Total media associated with cells: 634 Total labels associated with cell media: 343 |
Characters | |
Total characters: 210 Total characters with associated media: 1 Total characters with media with labels: 0 Total character states: 547 Total character states with associated media: 3 Total character states with media with labels:1 Total unordered/ordered characters:210/0 |
Currently Viewing:
MorphoBank Project 790
MorphoBank Project 790
- Creation Date:
12 September 2012 - Publication Date:
19 March 2014 - Media downloads: 1
- Matrix downloads: 76
This research
supported by
Authors' Institutions
- Natural History Museum, London
- Cornell University
- Florida International University
- Harvard University
- Stanford University
- University of Cambridge
- University of Wisconsin
- Queensland Museum
- Field Museum of Natural History
- Smithsonian Institution
- The Citadel: Military College of South Carolina
- Northern Virginia Community College
Members
member name | taxa | specimens | media | chars | character
| cell scorings (scored, NPA, "-") | cell
| rules | ||||||
Rudiger Bieler Project Administrator | 108 | 117 | 205 | 210 | 4 | 2 | 18995 (14631, 965, 3399) | 0 | 343 | 0 | ||||
Gonzalo Giribet Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 38 (25, 12, 1) | 0 | 0 | 0 | ||||
Dan Graf Character annotater | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Liz Harper Character annotater | 0 | 0 | 0 | 0 | 0 | 0 | 40 (26, 1, 13) | 0 | 0 | 0 | ||||
John Healy Character annotater | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Janeen Jones Observer | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Seth Kaufman | 0 | 0 | 0 | 0 | 0 | 0 | 1 (0, 0, 1) | 0 | 0 | 0 | ||||
Nikolaus Malchus Observer | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Nikolaus Malchus Observer | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Paula Mikkelsen Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 417 (298, 0, 119) | 0 | 0 | 0 | ||||
Prashant Sharma Character annotater | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Sid Staubach Observer | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Ellen Strong Character annotater | 0 | 0 | 0 | 0 | 0 | 0 | 33 (31, 0, 2) | 0 | 0 | 0 | ||||
John Taylor Character annotater | 0 | 0 | 0 | 0 | 0 | 0 | 132 (121, 0, 11) | 0 | 0 | 0 | ||||
Ilya Temkin Character annotater | 0 | 0 | 0 | 0 | 0 | 0 | 248 (88, 0, 160) | 0 | 0 | 0 | ||||
Thomas Waller Observer | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
John Zardus Character annotater | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M1796' (108 Taxa)
taxon | unscored cells |
scored cells |
no cell support |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] Leptochiton asellus Taxon name last Modified on 12/05/13 | 23 | 69 | 68 | 0 | 118 | 0 | 0 | 17 |
[2] Cryptoplax japonica Taxon name last Modified on 12/05/13 | 30 | 47 | 47 | 19 | 114 | 0 | 0 | 17 |
[3] Chaetopleura apiculata Taxon name last Modified on 12/05/13 | 24 | 68 | 68 | 0 | 118 | 0 | 0 | 17 |
[4] Puperita pupa Taxon name last Modified on 12/05/13 | 10 | 77 | 77 | 19 | 104 | 0 | 0 | 17 |
[5] Haliotis tuberculata Taxon name last Modified on 12/05/13 | 2 | 93 | 91 | 0 | 115 | 0 | 0 | 17 |
[6] Diodora graeca Taxon name last Modified on 12/05/13 | 10 | 73 | 71 | 19 | 108 | 0 | 0 | 17 |
[7] Crepidula fornicata Taxon name last Modified on 12/05/13 | 5 | 93 | 93 | 0 | 112 | 0 | 0 | 17 |
[8] Siphonaria pectinata Taxon name last Modified on 12/05/13 | 5 | 86 | 86 | 0 | 119 | 0 | 0 | 17 |
[9] Antalis entalis Taxon name last Modified on 12/05/13 | 7 | 84 | 81 | 0 | 119 | 0 | 0 | 17 |
[10] Laevipilina hyalina Taxon name last Modified on 12/05/13 | 17 | 85 | 84 | 0 | 108 | 0 | 0 | 17 |
[11] Solemya velum Taxon name last Modified on 12/05/13 | 1 | 118 | 88 | 0 | 91 | 26 | 13 | 17 |
[12] Huxleyia munita Taxon name last Modified on 12/05/13 | 78 | 80 | 80 | 27 | 25 | 0 | 0 | 17 |
[13] Nucinella giribeti Taxon name last Modified on 12/05/13 | 66 | 92 | 76 | 28 | 25 | 0 | 0 | 17 |
[14] Nucula sulcata Taxon name last Modified on 12/05/13 | 19 | 148 | 120 | 0 | 43 | 28 | 25 | 17 |
[15] Acila castrensis Taxon name last Modified on 12/05/13 | 11 | 152 | 128 | 0 | 47 | 22 | 17 | 17 |
[16] Clencharia abyssorum Taxon name last Modified on 12/05/13 | 19 | 119 | 94 | 20 | 52 | 21 | 23 | 17 |
[17] Scaeoleda caloundra Taxon name last Modified on 12/05/13 | 16 | 142 | 121 | 2 | 50 | 21 | 19 | 17 |
[18] Yoldia limatula Taxon name last Modified on 12/05/13 | 31 | 136 | 114 | 1 | 43 | 15 | 14 | 17 |
[19] Arca noae Taxon name last Modified on 12/05/13 | 21 | 168 | 134 | 1 | 20 | 22 | 20 | 17 |
[20] Barbatia barbata Taxon name last Modified on 12/05/13 | 15 | 175 | 143 | 0 | 20 | 28 | 26 | 17 |
[21] Arcopsis adamsi Taxon name last Modified on 12/05/13 | 26 | 145 | 122 | 19 | 20 | 20 | 15 | 17 |
[22] Glycymeris glycymeris Taxon name last Modified on 12/05/13 | 12 | 176 | 116 | 0 | 22 | 22 | 15 | 17 |
[23] Limopsis sp b Taxon name last Modified on 12/05/13 | 22 | 141 | 119 | 19 | 28 | 21 | 18 | 17 |
[24] Modiolus rumphii Taxon name last Modified on 12/05/13 | 20 | 167 | 124 | 0 | 23 | 18 | 11 | 17 |
[25] Mytilus edulis Taxon name last Modified on 12/05/13 | 7 | 180 | 130 | 0 | 23 | 20 | 10 | 17 |
[26] Pteria hirundo Taxon name last Modified on 12/05/13 | 20 | 173 | 141 | 0 | 17 | 3 | 0 | 17 |
[27] Isognomon alatus Taxon name last Modified on 12/05/13 | 20 | 167 | 132 | 0 | 24 | 31 | 11 | 17 |
[28] Malleus albus Taxon name last Modified on 12/05/13 | 21 | 146 | 116 | 19 | 24 | 16 | 2 | 17 |
[29] Pulvinites exempla Taxon name last Modified on 12/05/13 | 55 | 124 | 82 | 6 | 25 | 0 | 0 | 17 |
[30] Crassostrea virginica Taxon name last Modified on 12/05/13 | 16 | 169 | 138 | 0 | 25 | 15 | 1 | 17 |
[31] Hyotissa mcgintyi Taxon name last Modified on 12/05/13 | 65 | 111 | 82 | 19 | 15 | 3 | 0 | 17 |
[32] Pinna carnea Taxon name last Modified on 12/05/13 | 13 | 153 | 122 | 19 | 25 | 22 | 9 | 17 |
[33] Ctenoides scaber Taxon name last Modified on 12/05/13 | 15 | 171 | 170 | 0 | 24 | 0 | 0 | 17 |
[34] Lima lima Taxon name last Modified on 12/05/13 | 16 | 151 | 120 | 20 | 23 | 14 | 10 | 17 |
[35] Pecten maximus Taxon name last Modified on 12/05/13 | 23 | 166 | 132 | 0 | 21 | 22 | 11 | 17 |
[36] Parvamussium jeffreysii Taxon name last Modified on 12/05/13 | 16 | 165 | 134 | 0 | 29 | 24 | 9 | 17 |
[37] Spondylus ambiguus Taxon name last Modified on 12/05/13 | 30 | 154 | 111 | 0 | 26 | 7 | 0 | 17 |
[38] Plicatula sp Taxon name last Modified on 12/05/13 | 56 | 113 | 76 | 20 | 21 | 0 | 0 | 17 |
[39] Monia patelliformis Taxon name last Modified on 12/05/13 | 71 | 97 | 93 | 21 | 21 | 4 | 3 | 17 |
[40] Placuna placenta Taxon name last Modified on 12/05/13 | 33 | 148 | 110 | 3 | 27 | 1 | 0 | 17 |
[41] Dimya lima Taxon name last Modified on 12/05/13 | 61 | 106 | 78 | 20 | 23 | 0 | 0 | 17 |
[42] Neotrigonia lamarcki Taxon name last Modified on 12/05/13 | 18 | 171 | 142 | 0 | 21 | 16 | 4 | 17 |
[43] Velesunio ambiguus Taxon name last Modified on 12/05/13 | 29 | 162 | 161 | 0 | 19 | 0 | 0 | 17 |
[44] Aspatharia pfeifferiana Taxon name last Modified on 12/05/13 | 29 | 141 | 117 | 19 | 21 | 8 | 4 | 17 |
[45] Margaritifera margaritifera Taxon name last Modified on 12/05/13 | 25 | 142 | 137 | 19 | 24 | 4 | 3 | 17 |
[46] Unio pictorum Taxon name last Modified on 12/05/13 | 50 | 141 | 137 | 0 | 19 | 4 | 3 | 17 |
[47] Eucrassatella cumingi Taxon name last Modified on 12/05/13 | 13 | 174 | 169 | 0 | 23 | 4 | 2 | 17 |
[48] Astarte sulcata Taxon name last Modified on 12/05/13 | 14 | 174 | 172 | 0 | 22 | 3 | 3 | 17 |
[49] Cardita calyculata Taxon name last Modified on 12/05/13 | 17 | 166 | 163 | 0 | 27 | 2 | 2 | 17 |
[50] Frenamya elongata Taxon name last Modified on 12/05/13 | 14 | 156 | 151 | 1 | 39 | 2 | 3 | 17 |
[51] Lyonsia floridana Taxon name last Modified on 12/05/13 | 21 | 162 | 134 | 1 | 26 | 0 | 0 | 17 |
[52] Laternula elliptica Taxon name last Modified on 12/05/13 | 12 | 168 | 162 | 1 | 29 | 5 | 3 | 17 |
[53] Cochlodesma praetenue Taxon name last Modified on 12/05/13 | 24 | 143 | 136 | 19 | 24 | 5 | 2 | 17 |
[54] Thracia phaseolina Taxon name last Modified on 12/05/13 | 27 | 141 | 128 | 20 | 22 | 6 | 1 | 17 |
[55] Myochama anomioides Taxon name last Modified on 12/05/13 | 21 | 161 | 155 | 2 | 26 | 0 | 0 | 17 |
[56] Cleidothaerus albidus Taxon name last Modified on 12/05/13 | 27 | 142 | 109 | 19 | 22 | 0 | 0 | 17 |
[57] Haliris tenerrima Taxon name last Modified on 12/05/13 | 23 | 120 | 89 | 20 | 47 | 0 | 0 | 17 |
[58] Poromya illevis Taxon name last Modified on 12/05/13 | 26 | 110 | 96 | 0 | 74 | 0 | 0 | 17 |
[59] Bathyneaera demistriata Taxon name last Modified on 12/05/13 | 15 | 94 | 79 | 20 | 81 | 0 | 0 | 17 |
[60] Cavatidens omissa Taxon name last Modified on 12/05/13 | 50 | 129 | 121 | 0 | 31 | 4 | 1 | 17 |
[61] Codakia orbicularis Taxon name last Modified on 12/05/13 | 1 | 170 | 168 | 0 | 39 | 2 | 2 | 17 |
[62] Lucina pensylvanica Taxon name last Modified on 12/05/13 | 13 | 143 | 143 | 19 | 35 | 0 | 0 | 17 |
[63] Cycladicama cumingi Taxon name last Modified on 12/05/13 | 13 | 159 | 155 | 19 | 19 | 0 | 0 | 17 |
[64] Thyasira equalis Taxon name last Modified on 12/05/13 | 18 | 157 | 113 | 1 | 34 | 7 | 0 | 17 |
[65] Cyrenoida floridana Taxon name last Modified on 12/05/13 | 23 | 164 | 163 | 1 | 22 | 1 | 1 | 17 |
[66] Chama macerophylla Taxon name last Modified on 12/05/13 | 20 | 169 | 123 | 0 | 21 | 5 | 1 | 17 |
[67] Tellimya ferruginosa Taxon name last Modified on 12/05/13 | 76 | 94 | 94 | 21 | 19 | 0 | 0 | 17 |
[68] Lasaea adansoni Taxon name last Modified on 12/05/13 | 2 | 171 | 132 | 2 | 35 | 2 | 1 | 17 |
[69] Scintillona cryptozoica Taxon name last Modified on 12/05/13 | 18 | 143 | 102 | 22 | 27 | 0 | 0 | 17 |
[70] Mysella charcoti Taxon name last Modified on 12/05/13 | 20 | 137 | 102 | 21 | 32 | 2 | 0 | 17 |
[71] Bornia sebetia Taxon name last Modified on 12/05/13 | 54 | 118 | 115 | 21 | 17 | 2 | 2 | 17 |
[72] Panopea globosa Taxon name last Modified on 12/05/13 | 46 | 125 | 81 | 25 | 14 | 6 | 0 | 17 |
[73] Hiatella arctica Taxon name last Modified on 12/05/13 | 10 | 163 | 154 | 23 | 15 | 7 | 2 | 17 |
[74] Lamychaena hians Taxon name last Modified on 12/05/13 | 22 | 143 | 139 | 21 | 24 | 4 | 0 | 17 |
[75] Arctica islandica Taxon name last Modified on 12/05/13 | 8 | 179 | 174 | 4 | 20 | 4 | 2 | 17 |
[76] Trapezium sublaevigatum Taxon name last Modified on 12/05/13 | 27 | 165 | 131 | 0 | 19 | 0 | 0 | 17 |
[77] Glossus humanus Taxon name last Modified on 12/05/13 | 15 | 144 | 138 | 35 | 18 | 4 | 1 | 17 |
[78] Kelliella sp Taxon name last Modified on 12/05/13 | 52 | 94 | 92 | 20 | 44 | 0 | 0 | 17 |
[79] Calyptogena magnifica Taxon name last Modified on 12/05/13 | 62 | 105 | 93 | 28 | 15 | 2 | 0 | 17 |
[80] Cyamiomactra laminifera Taxon name last Modified on 12/05/13 | 129 | 53 | 53 | 19 | 9 | 0 | 0 | 17 |
[81] Gaimardia trapezina Taxon name last Modified on 12/05/13 | 69 | 98 | 69 | 28 | 15 | 0 | 0 | 17 |
[82] Corbicula fluminea Taxon name last Modified on 12/05/13 | 14 | 174 | 171 | 3 | 19 | 3 | 0 | 17 |
[83] Sphaerium nucleus Taxon name last Modified on 12/05/13 | 16 | 154 | 132 | 20 | 21 | 2 | 0 | 17 |
[84] Cerastoderma edule Taxon name last Modified on 12/05/13 | 24 | 165 | 162 | 4 | 18 | 2 | 0 | 17 |
[85] Fragum unedo Taxon name last Modified on 12/05/13 | 53 | 140 | 140 | 4 | 13 | 0 | 0 | 17 |
[86] Hemidonax pictus Taxon name last Modified on 12/05/13 | 25 | 167 | 135 | 0 | 18 | 2 | 0 | 17 |
[87] Venus verrucosa Taxon name last Modified on 12/05/13 | 22 | 150 | 148 | 19 | 19 | 1 | 0 | 17 |
[88] Chione elevata Taxon name last Modified on 12/05/13 | 16 | 171 | 169 | 5 | 18 | 2 | 2 | 17 |
[89] Gemma gemma Taxon name last Modified on 12/05/13 | 20 | 165 | 126 | 2 | 24 | 0 | 0 | 17 |
[90] Petricola lapicida Taxon name last Modified on 12/05/13 | 21 | 167 | 124 | 5 | 17 | 4 | 0 | 17 |
[91] Turtonia minuta Taxon name last Modified on 12/05/13 | 16 | 136 | 91 | 21 | 37 | 0 | 0 | 17 |
[92] Glauconome rugosa Taxon name last Modified on 12/05/13 | 24 | 168 | 166 | 1 | 17 | 2 | 0 | 17 |
[93] Scissula similis Taxon name last Modified on 12/05/13 | 18 | 163 | 153 | 0 | 30 | 9 | 0 | 17 |
[94] Macoma balthica Taxon name last Modified on 12/05/13 | 10 | 169 | 163 | 2 | 29 | 2 | 0 | 17 |
[95] Donax trunculus Taxon name last Modified on 12/05/13 | 21 | 166 | 148 | 0 | 24 | 14 | 9 | 17 |
[96] Asaphis deflorata Taxon name last Modified on 12/05/13 | 15 | 170 | 162 | 4 | 21 | 5 | 0 | 17 |
[97] Abra alba Taxon name last Modified on 12/05/13 | 21 | 157 | 123 | 4 | 28 | 5 | 0 | 17 |
[98] Tagelus plebeius Taxon name last Modified on 12/05/13 | 19 | 169 | 157 | 3 | 19 | 3 | 0 | 17 |
[99] Solen vaginoides Taxon name last Modified on 12/05/13 | 16 | 175 | 174 | 2 | 17 | 0 | 0 | 17 |
[100] Phaxas pellucidus Taxon name last Modified on 12/05/13 | 23 | 145 | 145 | 24 | 18 | 0 | 0 | 17 |
[101] Rangia cuneata Taxon name last Modified on 12/05/13 | 7 | 179 | 176 | 3 | 21 | 2 | 0 | 17 |
[102] Tresus capax Taxon name last Modified on 12/05/13 | 76 | 106 | 106 | 20 | 8 | 0 | 0 | 17 |
[103] Donacilla cornea Taxon name last Modified on 12/05/13 | 17 | 148 | 145 | 23 | 22 | 0 | 0 | 17 |
[104] Dreissena polymorpha Taxon name last Modified on 12/05/13 | 12 | 174 | 161 | 0 | 24 | 7 | 5 | 17 |
[105] Mya arenaria Taxon name last Modified on 12/05/13 | 5 | 173 | 170 | 12 | 20 | 2 | 0 | 17 |
[106] Notocorbula tunicata Taxon name last Modified on 12/05/13 | 45 | 144 | 144 | 5 | 16 | 0 | 0 | 17 |
[107] Barnea candida Taxon name last Modified on 12/05/13 | 24 | 159 | 126 | 0 | 27 | 2 | 0 | 17 |
[108] Teredo clappi Taxon name last Modified on 12/05/13 | 29 | 147 | 131 | 0 | 34 | 8 | 2 | 17 |
Project downloads
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 506 | |
Matrix downloads | 76 | BivAToL matrix (76 downloads); |
Project downloads | 415 | |
Document downloads | 14 | Additional information on sperm data for MorphoBank Project 790 - BivAToL (14 downloads); |
Media downloads | 1 | M148966 (1 download); |