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Gastropoda E Bivalvia.Key Mollusca 1 Quem são: Mexilhões, lulas, polvos, caramujos, lesmas etc. Cerca de 100.000 espécies viventes descritas e 70.000 espécies fósseis. 2 Aplacophora 3 Monoplacophora 4 Polyplacophora 5 Gastropoda 6 Bivalvia 7 Cephalopoda 8 Scaphopoda 9 LETTER RESEARCH , , yielded nearly identical topologies within Mollusca, except for relation-1 1–3,10 ships among basal gastropods and placements of the sea slug oxidase I c Gastropoda Pleurobranchaea and the bivalve Mytilus (Fig. 2 and Supplementary Pleistomollusca , Unnamed f , Scott R. Santos Fig. 5). High leaf stability scores for all OTUs (Supplementary Table4 3) . doi:10.1038/nature10382 10 a . Moreover, Mollusca was and strong support for most nodes suggest all OTUs were represented 3 Bivalvia Aculifer Institute of Zoology, Johannes Gutenberg by sufficient data to be reliably placed. Remarkably, branch lengths 4 . Unfortunately, because of varying 6 , Achim Meyer 3 were relatively uniform; cephalopods did not show long branches as Conchifera recovered most major lineages mono- 1–3,10 1 previously reported in analyses of 18S and 28S . Scaphopoda 00 MONTH 2011 | VOL 000 | NATURE | 1 Scaphopoda Bivalvia Monoplacophora Gastropoda Cephalopoda . Morphology has been interpreted to divide Neomeniomorpha Chaetodermomorpha Polyplacophora Conchifera 4 has been widely debated. Some workers have con- 1 4,9 All major lineages of Mollusca were monophyletic with strong , Diasoma and Cyrtosoma hypotheses allying bivalves to yielded similar results, namely that bivalves were not e 2 , Andrea B. Kohn 5 5 f support (bs 100%, pp 1.00). Importantly, there was strong3 Cephalopodaa Department of Biology and Centre for Geobiology, University of Bergen, P.O. Box 7800, NO- support at all deep nodes, although the node placing Scaphopoda Mollusca 2 Testaria Morphological and traditional molecular phylogenetic approaches Molecular investigations of molluscan phylogeny have relied Cyrtosoma Diasoma have failed to robustly reconstruct mollusc phylogeny. Notably, several recent phylogenomic studies (for example, refs 5cantly and advanced 11) our have signifi- understandingsequences of derived metazoan evolution from by genomeapproach, using and numerous transcriptome orthologous protein-coding data. genes Withtified can and this employed be in phylogeny iden- reconstruction. Many ofare these genes constitutively expressed andlimited can expressed be sequence easilygenes recovered tag are from (EST) even usually surveys.because informative of Additionally, their for conserved these nature inferring due to higher-level their phylogeny functional importance. hypothesis, allying scaphopods and cephalopods. Monoplacophora. scaphopods and gastropods to cephalopods, respectively. paraphyletic grade phyletic, but support at deeper nodesanalyses was of generally a weak. combined Subsequent data setand (18S, histone 28S, H3) 16S, cytochrome not recovered monophyleticApproximately Unbiased, AU, (a tests; Supplementary result Table 1)due possibly to significantly contaminated neomenioid supported sequences by sidered the presence of sclerites a synapomorphyuniting for a clade Polyplacophora Aculifera, (chitons;shells) and Aplacophora. which In contrast, Polyplacophora has have alternatively been both placed with scleritesMonoplacophora and Conchifera and Scaphopoda) (Bivalvia, in a Cephalopoda,the clade Gastropoda, called shelled Testaria uniting molluscs and have alsoanalyses offered of little 18S, resolution. 28S or Maximum both likelihood (ML) Conchifera into a cladebivalve/scaphopod (Diasoma) gastropod/cephalopod clade (Cyrtosoma) and a monophyletic and supportExpanding values on this at study, further mosttionship work between supported deep chitons a and sister-taxon nodes monoplacophorans rela- port (Serialia) at were but other deep sup- low. nodes was generally low received moderate support in ML (bs 5 72%) but strong support in interpretations of features as derived orsynapomorphies, and plesiomorphic, a often lack unclear character of homology, clear themorphology to ability of resolve such deep phylogenetic events is limited. primarily on nuclear ribosomal gene sequences (18S and 28S) BI (pp 5 0.98). A clade including Aplacophora and Polyplacophora & Kenneth M. Halanych Conchifera Polyplacophora Chaetodermomorpha Neomeniomorpha Bivalvia Scaphopoda Gastropoda Cephalopoda Monoplacophora 4 ,ora Aplacophora Neomeniomorpha 8 , Mathew R. Citarella was unequivocally supported (bs 5 100%, pp 5 1.00) and placed2 sister e .Whetherthese to Conchifera, consistent with the Aculifera hypothesis. Moreover, we , Adenopoda 7 .Manymolluscs 6 or morphological Macmillan Publishers Limited. All rights reserved found strong support (bs 5 100%, pp 5 0.99) for a sister relationship 1–3 Serialia Testaria , Bernhard Lieb Chaetodermomorpha , Hepagastralia hypothesis 3,5 b ©2011 between Neomeniomorpha and Chaetodermomorpha, supporting, Christiane Todt the 1–3,101 Department of Neuroscience, University of Florida, Gainesville, Florida 32611, USA. 5 Aplacophora hypothesis but contrary to previous molecular and .Additionally,withinConchifera, Aculifera hyletic group, Aplacophora )inacladewithamonophyletic Monoplacophora Polyplacophora Conchifera Polyplacophora Neomeniomorpha Chaetodermomorpha Neomeniomorpha Chaetodermomorpha Other Conchifera 4 omorphic state of Mollusca because , Aculifera hypothesis placing Aplacophora morphological studies. To evaluate alternatives to the Aculifera and Polyplacophora c abc d , Leonid L. Moroz , Serialia hypothesis allying Polyplacophora and Aplacophora hypotheses, we used AU tests (Supplementary Table2 5). d , Johanna T. Cannon 1 These tests rejected the Testaria hypothesis, which allies chitons The Whitney Laboratory for Marine Bioscience, University of Florida, 9505 Ocean Shore Blvd., St. Augustine, Florida 32080, USA. bs = 100, pp = 1.00 3 with the other shelled molluscs (P , 0.02) and placement of either Annelida bs = 100, pp > 0.98 llerweg 6, D-55099 Mainz, Germany. Leading hypotheses of molluscan phylogeny. a ¨ 95%) molluscan species. Thus we propose the node-based | . However, limited genomic resources spanning molluscan 5 , , have been unsuccessful at elucidating these relationships. aplacophoran taxon as sister to all other molluscs (both P 0.01). 4 Morphological disparity among the major lineages of Mollusca has Figure 4 | Deep molluscan phylogenyWith over 100,000 described extant species in eight major lineages, as inferred in the present study. Black Department of Biological Sciences, Auburn University, 101 Rouse Life Sciences, Auburn, Alabama 36849, USA. sister to Polyplacophora. University, Mu two lineages constitute a monop 5020 Bergen, Norway. Christoffer Schander are economically important as food and producerswhereas of others cause economic pearls damageas and pests, shells biofoulersand invasive species. Molluscs are alsostudy biomedically of important as brain organization, models learningof for and parasites. the memory Although as well shelledrecords as of molluscs any vectors have animal group, one evolutionarymolluscan relationships lineages among of have major been the elusive. best fossil prompted numerous conflicting phylogenetic hypothesesvermiform (Fig. Chaetodermomorpha 1). (also The Neomeniomorpha known (also known as as Solenogastres) traditionally Caudofoveata) have been considered and to represent theof their plesi ‘simple’ internal morphology and lack of shells data placing Neomeniomorpha basal. hypothesis placing Chaetodermomorpha basal. LETTER Phylogenomics reveals deep molluscan relationships Kevin M. Kocot Evolutionary relationships amongMollusca the have eight remainedimportance. Previous unresolved major investigations of despite molluscan lineages phylogeny, based theirprimarily of on nuclear diversity ribosomal gene and sequences Figure 1 1 diversity has prevented use ofuse a transcriptome and phylogenomic genome approach. data from Here allMonoplacophora) we major lineages (except andMollusca. recover Our a results stronglyplacing well-supported support Polyplacophora topology the (chitons AculiferaAplacophora for (worm-like hypothesis molluscs) asister-taxonrelationshipbetweenGastropodaandBivalviaissup- ported. This grouping has receivedmost little ( consideration and contains name Pleistomollusca. In light of these results, wetion examined the of evolu- morphological characterscephalization and and found shells support asMollusca. possibly for having advanced multiple origins within Mollusca is the second most speciose animal phylum Aculiferan monophyly supports interpretation of the Palaeozoic taxon circlesRecently, phylogenomic studies using dozens torepresenthave hundreds greatly of improved genes our understanding ofships deep animal relation- nodes with bs 5 100 and pp 5 1.00. Gray circles represent ‘Helminthochiton’ thraivensis as possessing features intermediate nodesNature, with bs 5 100 and07 pp de$ 0.98. Setembro The actual specimens ofdePolyschides 2011and between chitons and aplacophorans12, and interpretation of dorsal, Hanleya used in this study are shown. Photos are not to scale. A full-page serially arranged calcareous structures as a possible aculiferan synapo- version of this figure is presented in10 Supplementary Fig. 1. morphy13. Specifically, the chaetoderm Chaetoderma14 and some, but not all, neomenioids15 possess dorsal, serially repeated sclerite- the high degree of cephalization in gastropods and cephalopods has secreting regions during development. Notably, chiton valves are not recently been suggested to have evolved independently20.
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