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Phylogenetic Relationships of the Lower Caenogastropoda (Molluscs, Gastropoda, Architaenioglossa, Campaniloidea, Cerithioidea) A

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Phylogenetic Relationships of the Lower Caenogastropoda (Molluscs, Gastropoda, Architaenioglossa, Campaniloidea, Cerithioidea) A /yy// Zoológica Scripla, Vol. 27, No, 4, pp. 361-372, 1998 Elsevier Science Ltd Pergamon i;> 1998 The Noi'wegian Academy of Science and Letters All rights reserved. Printed in Great Britain PI I: 50300-3256(98)00017-8 " 0300-3256/98 S19.00 + 0.00 Phylogenetic relationships of the lower Caenogastropoda (Molluscs, Gastropoda, Architaenioglossa, Campaniloidea, Cerithioidea) as determined by partial 18S rDNA sequences M. G. HARASEWYCH*, S. LAURA ADAMKEWICZ, MATTHEW PLASSMEYER and PATRICK M. GILLEVET Accepted 9 June 1998 Harasevvych, M. G., Adainkcwicz, S. L. Plassmeyer, M. & Gillevet, P. M. 1998. Phylogenetic relationships of the lower Caenogastropoda (Mollusca. Gastropoda. Architaenioglossa. Cam- paniloidea, Cerithioidea) as delermined by partial I8S rDNA sequences.•Zool. Scr. 27: 361-372. Phylogenetic analyses of partial sequences spanning approximately 450 nucleotides near the 5' end of the IBS rDNA strongly support the monophyly of Apogastropoda and its constituent clades. Caenogastropoda and Heterobranchia. Representatives of the architaenioglossan groups Cyclo- phoroidea, Ampullariidae and Viviparidae invaiiably emerge within Caenogastropoda in all analy- ses. While the Cyclophoroidea and Ampullariidae are monophyletic, the varying position of Viviparidae in all outcomes contradicts its hypothesized sister group relationship with Ampul- lariidae, and thus the monophyly of Ampullarioidea. Because of the position of Viviparidae, Architaenioglossa does not emerge as a clade in any of our analyses. Campanile consistently emerges between Cyclophoroidea and Cerithioidea, or in a clade with Cyclophoroidea and Ampullariidae, a position not predicted by previous morphological studies. Maximum parsimony analyses of sequence data show Caenogastropoda to comprise a series of sequentially diverging higher taxa. However, maximum likelihood analyses as well as maximum parsimony analyses using only trans- versions divide Caenogastropoda into two clades, one containing the architaenioglossan laxa, Campaniloidea and Cerithioidea, the other containing the higher caenogastropod taxa included in Eucaenogastropoda (Haszprunar, 1988) [= Hypsogastropoda (Ponder & Lindberg 1997)]. Denser taxon sampling revealed insertions to be present in the 18S rDN/-\ gene of several caenogastropod taxa. Earlier reports {Harasewych ei al. 19976) of reduced sequence divergence levels in Caen- ogastropoda are shown (o be restricted to Hypsogastropoda. Based on a broader taxonomic sampling, divergence levels within Caenogastropoda are comparable to those found within Hetero- branchia. H 1998 The Norwegian Academy of Science and Letters M. G. Harasewych. Deparlnieni of inrerlebraie Zoology. National Miisetitn of Natiifal ¡iistniy. Sinitlisotiiati Institiitioti, Washitigton. DC 20560. U.S.A. S. Laura Aclaittkewicz. Departtiient of Biology. George Mason Utiiver.^ity. Fairja.x. VA 22030, U.S.A. Matthew Plasstneyer & Patricia M. Gillevet. in.stitnte for Bioscieitce. Bioittfonnatics. aitil Bio- teclttiology. George Mason Unicersity. Matiassas. VA 201 ¡0. U.S.A. Introduction Rosenberg el al. 1994, 1997; Harasewych el al. I991a,b). These recent studies converge on a broad outline of the The higher classification of the molltiscan class Gas- evolutionary history of Gastropoda, one significantly tropoda has remained static for much of the twentieth different from that formulated by Thiele and Wenz. Never- century, with most textbooks and reference works cur- theless, the placement and/or composition of a nuinber of rently in use incorporating an arrangement that was pro- key higher taxa remain unresolved or subject to conflicting posed by Thiele (1929-1931) and only slightly modified by interpretations. Among these are early radiations of land Wenz (1938-1944). Perhaps catalyzed by a reassessment and fresh-water taxa collectively termed the Arch- of prosobranch systematics begun by Golikov & Sta- itaenioglossa, and the superfamily Campaniloidea, rep- robogatov (1975), the past two decades have witnessed resented in the Recent fauna by a single, relict species. a renewed interest in re-evaluating relationships among Both groups figure prominently in the early history of higher gastropod taxa using large suites of anatomical, the Caenogastropoda, the superorder that contains the ultrastructural and molecular characters, and, more majority of living, shelled, marine gastropods. recently, cladistic methodology (e.g.. Salvini-Plawen 1980, Architaenioglossa has long been recognized as a ten- 1991; Salvini-Plawen & Haszprunar 1987; Haszprunar tative assemblage of terrestrial and fresh-water groups that 1988; Ponder & Lindberg 1996, 1997; Salvini-Plawen & could not be included elsewhere yet "bear little resem- Steiner 1996; Healy 1988, 1996; Tillier el al. 1992, 1994; blance to one another" (Thiele 1929). The group originally consisted of four families, the terrestrial Cyclophoridae, ' Corresponding author. and the fresh-water Viviparidae, Ampullariidae, and Lavi- 361 Zoológica Scripta 27 362 M. G. Harasewych et al. geriidae (Thiele 1929; Wenz 1943), although, the Lavi- phylogenetic analyses of Salvini-Plawen & Steiner (1996) geriidae were subsequently transferred to the Cerithioidea (Fig. ID). However, the analyses of Ponder & Lindberg (Morrison 1954). Most traditional classihcations have (1996, 1997) (Fig. IE and IF, respectively) indicate that placed the ai'chitaenioglossan families at the base of Caen- Architaenioglossa emerge within Caenogastropoda, as sis- ogastropoda or its paraphyletic component, Meso- ter group to all remaining caenogastropod taxa. gastropoda {e.g., Thiele 1929; Wenz 1943; Taylor & Sohl The monophyly of Architaenioglossa has been ques- 1962; Boss 1982; Ponder & Waren 1988; Vaught 1989). tioned by many recent authors {e.g., Sitnikova & Sta- Architaenioglossa was elevated to ordinal rank (Golikov robogatov 1982; Haszprunar 1988; Salvini-Plawen & & Starobogatov 1975; Ponder & Waren 1988) to include Steiner 1996; Ponder & Lindberg 1997), who noted the the superfamilies Cyclophoi'oidea and AmpuUarioidea (as absence of well defined synapomorphies to unite the Cyl- Viviparioidea), the latter containing the families Ampul- cophoroidea and AmpuUarioidea. Haszprunar (1988: 415) lariidae (as Pilidae) and Viviparidae. Golikov & Sta- further observed that the monophyly of AmpuUarioidea is robogatov (1975) (Fig. I A) regarded Architaenioglossa to not well supported, since apart from sharing a specialized constitute a lineage not closely related to taxa included osphradium, the Ampullariidae and Viviparidae are in Caenogastropoda by most authors. Haszprunar (1988) "remarkably dissimilar" in their nervous, respiratory and (Fig. IC) placed Architaenioglossa outside the Caen- excretory systems. Healy (1996) reported that Cyclo- ogastropoda, as sister group to Apogastropoda [ = Caen- phoroidea, AmpuUarioidea and Cerithioidea share a series ogastropoda -I- Heterobranchia, as redefined by Ponder of specialized eusperm and parasperm features, and form & Lind berg, 1997]. This position was reaffirmed by the a distinct group within Caenogastropoda. Monoplacophora ] Palemones Docoglossa Docoglossa SCUTIBRANCHIA 1 PLEUROTOMARIIONES SCUTIBRANCHIA TROCHiFORMII OPISTHOBRANCHIA BUCCINIFORMII PULUONATA Tnphoridao Triphoridae CONIFORME I PytamidsUomorpha CERITHIOIDËI Art^itectonicoideaa _]. RISSOOIDEI NERITOMOHPHA\ CYCLOPHOROIDEI CERITHIIDAE LITTORINOIDEI TROCHIONES MUfllCOIDEA I CERITHIOMORPHA Seguenziidae CONOIDEA CASSIDOIDEI XENOPHOROIDEI TURBINIWORPHA\ VIVIPAROIDEI CYCLOPHOROIOEA])EA] Vah/atoidei AWPULLARIOIDEA | PALUDINIMOPPHA Neomphaloidei CYPRAEOIDEI ^ Valvaloidaa NERITOPSIFORMII LITTORINIDAE Archileclonicitormii Z3 Pyramidelliones OPISTHOBRANCHIA Z] BULLIONES TONNOIOEA UTTORINIMORPHA ^ PULMONATA ID SIPHONARIIONES Seguenziidae GOLIKOV & GOLIKOV & STAROBOGATOV, 1975 B A STAROBOGATOV. 1988 Docoglossa Monoplacophora Neo)0pelopsidae Cephalopoda Cocculinilormia Docoglossa NERITOPSINA Cocculinjformia Melanodrymildae NERITOPSINA VENT TAXA Naomphalidae :] VETIGASTROPODA VETIGASTROPODA Uelanodryfniid; Seguenziina Neomphalidas CYCLOPHOROIDEAl D' ARCHITAENIOGLOSSA ARCHITAENIOGLOSSA AMPULLARIOIDEA J CAENOGASTROPODA CAENOGASTROPODA CAMPANILOIDEA CAMPANILOIDEA Valvatoidaa Valvaioidea Archiieclonicoidea Afchitectonicoidea Rissoelloidea Rissoelloidea z m HETEROBRANCHIA 0 :D Glacidorboidea P^ramldeltoidea 1 O Pyramidelloidea PULMONATA EUTHVNEURA OPISTHOBRANCHIA SALVINI-PLAWEN & HASZPRUNAR, 1988 STEINER, 1996 Polyplacophora Polyplacophora Monoplacophota Monoplacophora Cephalopoda Cephalopoda Docoglossa Docoglossa VETIGASTROPODA NERITOPSINA NERITOPSINA VENT TAXA Vent Taxa VETIGASTROPODA CYCLOPHOROIDEA" CYCLOPHORIDEA ~ AMPULLARIOIDEA AMPULLARIOIDEA CERITHIIDAE CERITHIIDAE Triphoridae CAMPANILIDAE CAMPANILOIDEA Triphoridae LITTORINIDAE rC LITTORINIDAE TONNOIOEA RANELLIDAE MURICOIDEA MURICOIDEA CONOIDEA _ HI CONOIDEA _ Valvaioidea Valvaioidea Architectonicoidea Archiledonicoidea OPISTHOBRANCHIA il OPISTHOBRANCHIA PULMONATA ^ PULMONATA _ PONDER & LINDBERG, 1996 PONDER & LINDBERG, 1997 Fig. !. Recent morphology-based hypotheses of gastropod evolution. Nomenclature and taxon rank have in some instances been modified to facilitate comparisons. Taxa listed in UPPER CASE are represented in the present study.•A. Golikov & Starobogatov (1975: hg. 6).•B. Golikov & Staroboeatov (1988: composite of figures 2, 3, 6. 9, 16. 17 and 27).•C. Haszprunar (1988: ñg. 5).•D. Salvini-Plawen & Steiner (1996; composite of figures 2'^6 and 2.7).•E. Ponder & Lmdberg (1996: fig. 11.3 D).•F. Ponder & Lindberg (1997: fig. 5).
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