Molecular phylogeny of some indo-pacific genera in the subfamily Turrinae H. Adams and A. Adams, 1853 (1838) (Gastropoda: Neogastropoda)

We have carried out a phylogenetic analysis of a group of Indo-Pacific species in the subfamily Turrinae (Swainson, 1840) using 128 mitoehondrial ribosomal RNA gene sequcnces Most of the species analyzed are convenlionally assigned to one of there Lophiotoma Casey, 1904. The inolecular anaysis revealed that while the species of Turrin and Gemmula analyzed in this study comprise monophyletie groups. the species presently assigned to Lopholoama definitely do not conslitute a monophylelic assemblage and can be separated into two very distinctive groups of species lased on the molecular analysis. The species presoutly designoted as Laphiationa tayabasensin Olivera, 2004. Lophiotoma panglaoensis Olivera, 2004. Lophiotoma indica (ricling, 1798) and Lophiotoma (Unedogemmula) uncdo (Kiener, 1839 in 1834-50) by molecular criteria, and are clearly widely separated from Turris, Genmmula or Lophiotoma (as redefined). We propose that Unedogemmula (MacNeil, 1960) be recognized as a full genus: Uncedogemmula unedo (Kiener, 1839 in 1834-80) in the lype spectes, and the specics above are transferred from Lophiotoma to Unedogemmula.

THK NAUTILUS 121(3):13l-138, 2007 Page 131 Molecular phylogeny of some Indo-Pacific genera in the subfamily Turrinae H. Adams and A. Adams, 1853 (1838) (Gastropoda: Neogastropoda) Fraut'isco M. Ileralde III National Institute of Molecular Biology and Biotechnology and Marine* Science Institute University of the Philippines Diliman, Quezon City PHILIPPINES Cisela P, Conception Marino Seipiiec Institute University of the Philippines Diliman, Quezon Citv PJJIMl'PJNES ‘ Maren Walkins john-Psuil Owtnby Pmdip K. Bandyopadhyny Department of Biology University of Utah Stilt Lake CJity, UT 84112 USA Baldomero M. Olivcra1 Department of Biolojiy University of Utah Sail Uke City, UT S41I2 USA oliveni©l>iologyaitah.cdu Amcurfina D. Santos National Institute of Molecular Biology and Biotechnology University of the Philippines Diliman, Quezon City I'lm.nTiNrcs ABSTRACT Wo have carried onl a phylogenetic analysis of a group of Iiulo- Pacific species in the stihlamily Tunlnae (Swainson, 1840) us­ing I2S initwlionclrial rihosomal UNA none sequences, Most of the species analyned tire conventionally assigned to one of three gcnci'iu Turrlx Boding. IV9S, Cmnmda WeiiikaulY, I&7G or L<r))hU>tomn Casey, !$)()-(, The molecular analysis revealed that while the spedcs oi Tutrix imd Cvmmulti analyzed in this study tsjinpri.se monophyletic groups, the species presently assigned to Luvliiohmn definitely do nol constitute a monopnylclic as­semblage and cun lie separated into two very distinctive groups of spccies bused on the molecular analysis. Thu species pres­ently designated as Liphhit/ww totjabamtiabi Olivera, 2004, /lophiofonur lumglaornxh Olivcra. 2(KV1, Loph'mhimn indict* (Riiding, im) and Lophiatoma bhrnj/i Olivera. 2004, ;ir<- i<>- latcd to h>/>hi0i<>»i(i (Umviojymmufn) nncdo (Kiener, liStlVJ in 1834--S0) by molecular criteria, anil arc clearly widely sepa­rated from Tunis, G^imnnln or Lctphloloma (as redefined). We propose that Uiittdoipmmulti (MaeNeil, I960) be recognized as a frill genus; Uncdoi'cntmiilti unrtlo (Kiener, 183S in 15S34-W)) is vlui lype species, and the species above ;ire transferred from Lophlotanm lo Uiiakgciuiittda. INTRODUCTION Venomous gastropods comprise throe groups: the cono snails, the auger s noils or terebrids and this "turrids", classically included in a single family, Tunldae (H. Ad­ams and A. Adams, 1853): these families arc generally assigned to the SUpcrfaniily Omoklen (Tender and 1 Author for correspondence Waren, 1988; Taylor, Kuntor, and Sysoev, 1993), It has been apparent for some time that the "tiirrids" comprise the largest species group in tlie supcrfamily (see for ex­ample, Powell, 1966); however. the groundbreaking work of Bouchet and co-workers (Bouchet et al, 2002; 2004) in New Caledonia, has provided evidence that *90% of Conoideart biodiversity probably resides in the "lurrids" (broadly defined). TurridjT are problematic at all kvels; not only are they a megadiverse group (>10,000 species) with a large fraction of species that remain undescribcd, but the phy­logenetic relationships vvithin the group are poorly un­derstood. The number of different "ttirrid" genera (hat have been proposed is >600; although in traditional mol- hiscan taxonomic work all turrids had been assigned to the fan-lily Turridae, in most of the more recent system­atic treatments, the group hits been split into 3-6 differ­ent families (Taylor ct al., 1993; Bouchet and Rocroi, 2005). However, some standard taxonomic treatments retain the old nomenclature (see for example Kolia, 109S). To complement our ongoing stndy of turrid venoms (see for example Watkins et al., 200G) we initiated a,study of "turrid" molecular phylogeny; the first results at the generic level are reported below. Tlie genus Tunis Uiiding, 1798, is the nominate genus of the family Tur- ridae, with Tunis babijfoniti {Linnaeus, 1758) as the type specics. This is an exclusively Indo-paeifie genus. How­ever, there has been inconsistency in the definition of those genera, which are traditionally grouped together with Turns iu tllo subfamily Turrinae. Two olher groups largely from live Indo-Pacific, Lophiotomn UVmkuilT. Ih75, and Gvmmuhi Cuscy, 1904, are included in (hePiige 132 THE NAUTILUS, Vol. 121, No, ii subfamily by most workers; in the western Atlantic and eastern Pacific, the genus Polys lira Woodring, 1928, is also thought lo have a close affinity to Trttris (Ihe type species for these genera are Lophiotoma (tcuid (Peny, 1811), Gemmula hlntkitmu (Buriy, I95S) suid Polystira alhkh (Perry, 18J1)). Additionally, u number of other groups {such its Titrridnipa (Hedluy, 1022)} are regarded as turrine by most workers, A major motivation for these studies is io investigate thw toxin genes expressed iu llie venom ducts of couokloiin gastropods. Among the different groups of turrids, we lwve initially concentrated on studying the gens products expressed in the venom ducts of species in the subfamily Turrinac, since they are larger and more easily collected than are most other turrid groups. Wc hope lo correlate tlie gene families expressed in venom ducLs willi the molecular phytogeny of the species ana­lyzed Thus, the molecular analysis reported below has focused on larger Indo-Paeitic species in the subfamily Tun'inae, e.g. Turrii, Lstphiotomri and Gemmula spp. The data that we present below demonstrates that two groups of spceies presently assigned to the genus Lojthiotaim, which appear to be closely related by shell morphology, prove to he unexpectedly divergent when evaluated by molecular criteria and need lo be placed in different genera. MATERIALS AND METHODS Specimen Collection: Species analysed in this Study, shown in Table 3, were mostly collected by commercial shell collectors in the Philippines, except for Pohjslim alblrfu (t'erry, 1811), a generous gift of Drs. Estuardo Lopez Vera and Ed Ilcimer, and Lophiohmc ccritlufor- mix (Poivcll, U)fi4), which was collected in Oahu, Hawaii, Specimens of cach were preserved either in RNAlaler® (Ambiou Inc., Tx) or 95% ethanol and DNA extracted as described below. In most cases, llie digestive gland was used as the source of DNA; however, for alcohol pre­served specimen,'! where the shell had not been craekod, the* digestive gland was often degraded, and DNA was extracted from foot tissue. Identification und sequencing of clones encoding 12S mitochondrial rHNA gene segments: Genomic DNA was prepared from tissue (~20 mg) from each turrid species using the Gcntra PUKE GENE DNA isolation kil (Gcn- tra Systems, Minneapolis, MN} according to the manu­facturer's standard protocol. Genomic DNA from each species (-*10ng) whs used as a template for polymerase chain reaction (PCtt) with oligonucleotides correspond­ing to 12S-I (S' TGC GAG CAG YCG GGG TTA ) and I2S-11I (5' AGA GYG HOG GGG GAT GTG T) mito­chondrial r|iNA segments (Oliverio and Mariottini, 2001), The 5' and 3' primers included adapters GOAGA- OAU and GGGAAACU respectively for annealing to the cloning vector pNEB206A. The PCIIl cycling profiles were as follows: initial denatunltioil (95°C, 60s); followed by -10 cycles of denaturation (05"C. 20s); annealing (55r‘C, 20s) ami extension (72°C, 30s), Tlit* PGR products were Table 1, IJ-jt of species analyzed in tins study. Locality Spccii fjiphkitoimr acuta (Perry, ISH) Lophioiomti biseuia' Olivera, 2004 Ij)phiotvnUi Lcrithifor'iith (Powell, 19G4) Txiphiotmna ([.iirnrtrok, 1822) Lophioioma indica' (Ruding, 1708) l*t>f>hiou»utt jickulii (Winnkauii; 1S75) Lophiolaiim ohmgoensis Olivers, 2002 I.ophiuioMrL Olivers, 2004 Lophiotoma pohjtropa (Ilelbling, 1770) f.(i)>hi(iloiria InijohrlsiMaU* Olivt.ru, 2004 Lnphiotoma uneth' (Kicner. IS 3D in 1834-80) Gemmula .hjcwwi (Reeve, 1843) Gemmula diomodea Powell, 1964 Gvmiimltr rtixarh Shikama mil I Hayashi, UJ77 Gemmula Iteajmii Olivera, 2000 Canmnla sugodatttls Olivera, 200,5 Tntti/s jjdnu»tgll (Reeve, 1843) Turns grandis (Gray, 1833) Turris nommn clavkhon I Olivera, 2000 Tunis babyUmht (Linnaeus. 1758) Turrit spectablllt (Reeve, 1843) Turrii totiphjtfix Olivera, 2000 PohjHlfyt all/Ida (Periy, 1811) Driflia regina (Ilabe and Murakami, 1970) Buemvista, Mn.mi;lnque, Philippines BuUUH^s, Philippines Oahu, Hawaii Guvoy, Olango Island, Philippines Aiijjway is. Dipolog, Philippines Cawoy, Qlnnjro Island, Philippine:;, Cawoy, Olango Island, Philippines Paugko (s. Boliol, Philippines Halaan, Luzon, Philippines Segod, Cebu, Philippines Piinglao Is. Bohol, Philippines Batanj»as Philippines Sugod, Cebu, Philippine* Sogod, Cebu, Philippines Sogod, Cebu. Philippines Sogod, Cebu, Philippines Cawoy, Olango Island, Philippines Sogod, Cebu, Philippines So^od, Cebv, Philippines Cawoy, Oianj'o Island, Philippines Cawoy, Oluii^O I.d:uid, Philippines Cawoy, Olango Island, Philippines Bay of Campeche, Mcxk-o Fangluo Is., Bohol, Philippines * These species are proposed to be transferred from Lophiotomu to UHwfajifminma (see toxt)- purifled using the PureLink PCK Produet Purification Kit (Invitrogcn Life Technologies, Qulsbad, Califomia) following the manufacturers suggested protocol. The eluted DNA fragments were digested with uracil specific excision reagent, annealed to pNEB206A vector (USER™ Friendly Cloning kit, New England lJioLabs. Inc., Beverly, Massachusetts) and the resulting products trails formed into competent DI15a cells (Sambrook andF. M. Ileralde III et al., 2007 Page 133 Russell. 2001). Plasmid DNAs wore isolated from ampi- cillin resistant colonics and the nodeie acid sequcncra of the inserts determined using ABT DNA sequencer with ABI Uig Dye ehcmistiy (Foster City, CA), DNA se­quences have been submitted to GenEank and the accession numbers are: EF467333, EF4673334. EF467335, EF467336, EF467337, EF467338. EF467339, EF4S7340, EF467341, EF467342, EF467343, EF467344, EF467345. EF467346. EF467347, EF46734S, EF467349, EF4673S0, EF40735I, EF467352, EF467353. EF467354, EF467355, and EF4B7356. Sequence Analysis; Nucleic acid sequences (the long­est of which hau593 nucleo tides) were aligned manually using MEGA version 3.1 (Kumar, 2004), One tree was creatcd from Iwo independent runs vising the software program MrBuyes (Iluelsenbeck, 2001; Ronquist, 2003). 5.000.000 trees were in tide in cach run, 50,000 of which were saved. Two hundred and fifty of each of those 50,000 were also discarded as burn-in. Each run hud four chains (one cold and three heated), The two independent runs were combined Into a single tree where branches were preserved if they were found in 70% or more of thosu trees not discarded, The standard deviation after 5.000.000 generations was 2.401 x 10 A general time reversible (GTIl) model was used, with the rate variation of some sites kept invariable and the remaining rules drawn from a gamma distribution. The other tree was created using the software program PHVML (Guindon, 2003). A thousand trees were obtained using non- parametric bootstrap analysis and combined into ji single tree where branches were preserved if they were found in 70% or more of the given trees. A GTR model was used, with the base frequency estimates found empiri­cally and the proportion of invariable sites estimated. Four substitution rates were used, wilh the gumma dis­tribution parameter estimated, RESULTS AND DISCUSSION PCR Amplification of 12S Sequencing: The se­quences of 12S rDNA from 23 species in the subfamily Turrinae (see Table I) were obtained as described above. The 12S sequence of a Drilli/t spedes. Dridkt ref'iun (Ilube and Murakami, 1970), was used as the outgroup for the phylogenetic analysis. The sequences obtained are shown in Table 2; these were aligned for maximal overlap. Phylogenetic Analysis: A phylogenetic tree, shown in Figure 1, was constructed as described under Meth­ods. The species that are presently assigned to two of the major turrino genera, Tunis and Ccmiiitifri, appear as monophyletic eludes in the phylogenetic tree obtained through Bayesian methods, However, the Lophiotoma species analyzed dearly split into two distinct, well- separated groups. Thus, the spedes presently assigned to Jjophiotomn analyzed in this study do not appear to constitute a mouophylfctie assemblage. A large separation is found between two groups of Lophiotoma species; one group includes Lophiotoma uncilo (Kioner, 1839 in 1834-80), Lophiotoma laijabrtisen&is Olivera, 2004, Lophiotonw prrngfonfinsls Olivers, 2004, Lophiotom/i indica (Rtiding, 1798), and Lvphlotonm bisaya Olivera. 2004. These spe­cies ;tppear to be much more divergent from the Tuiris and Gainmula branches than the other group of Lophiotoma, which includes the type species of Lophiotoma, Tjaphlofoma acuta (Perry, 1811),- the latter comprises two sub-branches, one branch including spe­cies such iis Lophwtojim tingulifera (J^imarck, 1522), assigned by many systematic to the .subgenus Xcntir'o- Uirria Iredale, 1929, which is regarded as a separate ge­nus hy some workers (Powell, 1966). Generic Classification and Nomenclature; The un­expected phylogenetic separation between two groups of species conventionally assigned to the genus Lophiotoma, makes the present assignment of these spe­cies into the conventional Indo-Paeifie Unrine genera, Turriff, Lophiotoma, and Gcmmtila inconsistent with the phylogenetic tree shown in Figure 2. One potential so­lution would be to lump these Indo-Paciiie genera to­gether under one genus, Turrit, and use subgeneric des­ignations for each large clade of species (this might be called the "Coitus alternative"; the major group oflndo- Paeilie Turrinae comprise a phylogenetic branch that does not appear to be more divergent by molecular cri­teria Ilian is the divergence within the species presently assigned to the genus Count;; Expiritu, 2001). Although this alternative may have some merit, the substantial lit­erature referring to species in the traditional genera Tur­rit, Gcmnuth, and Lo])hioUniui (including a significant paleontological component of the research literature) would make this A radical (and probably impractical) al­ternative. Because I^ophiotima acuta is the designated type spe­cies for the genus Lop/iiotniun, the species presently in Isiphiolonia that are in the branch not including L, aculd require a new generic designation. There arc two ge- neric/subgenriric designations potentially available for the group. One is a name proposed originally by Powell, Lophfoturtii Powell, 19fi4; Powell envisioned Lophlotur- ri.v as a genus allied to Utpldotomn with Lophiotomi indica (Rodirtg, 1798) as type, hiphiotnrris was set up specifically for forms that have blunt paucispiraf proto- conchs; since most of the species in this chide have po- lygyrote (multispiral) protoeouehs, U>phlotuni& does not seem to be an appropriate taxonomic designation (see Powell, 1964, for a discussion of differences in proto­couch motphology). The other available generic name for this group of spedes (which would have priority) is UnedojiCHimiila MacNeil I960: as originally proposed, Uncdogemtnnla was a separate genus, with Unrlo&mmuln uncrto (Kiener, 1S39 in 1K34-fiO) as type. However, Powell relegated Unnlogpmmitla to be n subgenus of Gvmnmla. Subse-Table 2. Alignment of 12S rDMA sequences. 1. Lophiotoim ceritlttfannis. 2. Loptiiotonm cin«uUfer<t. 3. Lophintoma ohngpcmh- 4. Loplihttmw layabasenslt. 5, Lojj/iifitaMW to nimbi. 6. Lopfiiofujiia puiijtn/jm. 7. LophtolinmtjickeliL 8, Lophlotomapmtgltioemte. 9. Luphivtoma indica. 10. Lopiiiofaitiabhaija. 11, Turrit loliphyfllx. 12. Turrhspectaliilis. 13. Tunis babijlonia. 14. Tur/is gruiiflis. 15. Turris noniuincfavidsoni. 16. Turns "ariumsi'i. 17. Genmnh sogtxlensis. 18. Gennnuht diomedea. 19. Ccmmii/rr spariosu. 20. Lophlatatm atvla. 21. Ganwtuhi jkofoni. 22. Gcmamla ntsario. 23. 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Hail : 2 OF. M. Heralde Til ct al.. 2007 Page 135 Lophiotoma pofytrvpa C*mm‘>l‘>r0sori<> Figure 1, Pliylogcm.-lic tree of specks listed in Table L, A tree buSiid on 12S i'DNA sequCMieeS was constructed (sou Mcdi<xLi). Species from three Indo-Fndfu: families in the subfamily Tuniiirte wire analyzed {Tunis. Gt'nwiula, hsphh>h»n<t). In uddilirm. one Atlantic species, TohjMira tribute and a spceics in the subfamily Drillimur (or Clavirue), DriUki I't^iun wm' included. N«la tlmt the species ot Lo))kit)loma arc split into two widely separated groups. As discussed in the text, specie's marked by an asterisk are prnpost'ti to be* movcU to iinodw genus. Lojihiotoma jfckclii and Turns gmntfit arc non-stiinilard taxomomic assignments for tho iigmvd specimens-: llu-sc arc widely regarded :>s synonymous to Lophiotoma matta und 7)/rri> cmjio. respectively (sco Powell, UXvJ), However, the molecular data clearly separates I^ophlotoind acula and Lophioloma jickrlii, und tw h-pe of Tunis crhtpa is suffieienliy dkergt-nt from the form shown dint using Turrit grnndis seems a preferable inune for this species. qiwnHy, Klllmm (1983) suggested that based on general shell morphology, Vnedogeuunula seems muck closer to species traditionally assigned to Lophiotoma. and that it should more appropriately be regarded us a suljgcnus of Lophh>to>na\ this suggestion has been adopted in most of the recent taxonomic treatments of turrine genera. The molecular results reported above demonstrate Hint Vuedoff-'tnmula uncdo is indeed closely rotated to some of tlie spccies most taxonomists presently include in Lophiotoma, (such us Lophiotoma Indira. Lophiotoma hhnijfi. ami Lophiotoma tciijabasemis). Furthermore, the molecular analysis clearly shows that there is no justifi­cation for designating Unedogemmtda as a submenus of either Gemmula or Lophioloma, since Unedoffnmmda unedo is in a very divergent branch of tlie phylogenetic tree. Thus, our results support the origimd designation of Uncdogcmmulct as u full genus, although tlie spceies comprising the genus need to be somewhat redefined. Unetfogemimiln uncdo is the type species, and tlie larger, strongly maculated forms previously assigned to Lojjhiotomn (such as Lophiotoma indica, Lophiotoma iatjabasensia, and Lophiotoma bi-taya) are transferred to Unedogeimnid/t from Lophiotonw. A recent analysis of Philippine forms related to these .specie,s. elarilymt; the relationships between these forms and the Unadogcm- multi uncdo group, was recently published (Olivera,Page 136 THIS NAUTILUS, Vol. 121, No, 3 Polystira albida ' Driliia rugius 98 96 Vnedogemmula pangiaaensis **%► 7Q|■ Vnedogemmulatayabasemis i- Unti ^ 84 95 91 100 100 jen Unahgemmtfawudo ■ Vnedogemmula indica Unedogemmula bisaya Tunis normandavidsoni -* Tunis garnottsii . Turris grandis Tunis bobylonia 86 83 Tunis spectabilis 1001- Turris totiphyllis Gemmula diomedea - Gemmula sogodensis ^ ■i- - Gemmula speciosa c-: Gemmula lisajoni - Gemmula rosario 100 c 100 86 Lophiotoma polytropa Lophiotoma jickeUi - Lophiotoma acuta lOOf---' Lophiotoma olangoensis -T" 1001------- 100 ij-------- >- Loi 0.1 expected substitutions per site Lophiotoma cerithiformis Lophiotoma cingutifera 4^j£j£$$ss»- Figure 2. The plyliigenetic tree la Figure I is rc-platti'd, except tluu llie confidence limits are included (calculated after 10 million generations), Note that all tin: major groups are highly supported; as is explained in the discussion, one of the groups formerly assigned to Lophiotoma is now regarded as a full geims, UnedogcmniuJa, with Unecbj^tnnmlti nuedo as type species; ilrns, in this figure, tin:jc spccies (still labeled Lnphlatmnu in Figure 1) are designated is Unedtwemninkt spp. 2004); it seems likely that all of the species level tux a, treated in that work are properly placed in the genus Unedoj'firrwmla, as redefined below, although this should be verified by obtaining molecular data for those species. Genu? Unedogcmimln Muc-Neil, I960 Description; Shell large, SO-lOomm, fusifonn with tall spire, long, straight, anterior canal unnolched. Pro- loconch variable, from blunt paurispirnl to lindUspiral, often with the transitional part of the larval shell deco­rated with brephie uxials or a.\iallv co.state whorls. Sinus is peripheral, deep, and narrow at the (emiiiuUion of the sinus rib. Uemarksi The genus has shell morphology with strong similarities to lA>j)hiotorna: most species havo a smooth peripheral keel but, in some sperics, there are distinct peripheral granulations. In otbev species, these granula­tions continue to the body whorl, hut these tend toward obsolescence in most forms. In contrast to Lophiatoinn, the peripheral lewd does not consist of two raised cords at the edges with a depressed area in the center; rather, inF. M. llrnlde III et III, 2007 Fugc 137 some forms, there is a single smooth rib exhibiting a variable level of peripheral granulation. In order to test the veracity of the proposals based on tlie Bayesian analysis and outlined above, a second ap­proach to the phylogenetic analysis of the sequences was also carried out. A Maximum Likelihood method was employed as described under Methods. The results of this analysis are shown in Figure 3. The separation of the speeies described above originally in Jjiphlotoma iuto two distinct groups is strongly supported using this analy­sis, Thus, both methods support raising Umdognnmula to a lull genus, as described above. Hie analyses differ, however, in the results with Gemmula; the maximum likelihood method does not group all of the species ana­lyzed into a single rrionophylctic chide, but into two groups of species. Thus, given this discrepancy between the two methods, the monophyly of Gantntuh clcarly requires further investigation, A brief summary of a proposed revision of Tndo-Padfie genera in the subfamily Turrinae is given in Table UL The eludogram in Figures 2 and 3 give support to Turri-y and Unc'ilogtffitiiiula. However, Lophioloma (redefined to exclude the -species transferred to Vnodogvmmuh) has two branches, both strongly supported: the conjoining of the branches lias loss than y0% support in the Bayesian analysis, and is not supported above the cutoff level 97 87 ■ I. Drillia regius Polystira albida - Unedugemmula panglaoensis - Unedoganmula tayabasensis Unedogemmula uitedo - ' - Unedogemmula indica 84 72 85 99 97 Unedogemmula bisaya --- Tunis normanda vidsoni - .-i ' Turns gamonsii - Turris grandis ■ Turris babylonia • Turris spectabilis TooL' 84 ■ Turris totiphy/lis Gemmula diomedea - Gemmula sogodemis - Gemmula speciosa 1001---------' Gemmula lisajoni I ■ ■■ - - Gemmula rosario 95 Lophiotoma polytropa Lophiotoma jickeffi ■ ■ Lophiotoma acuta 97 m 98 > Lophiotoma olangoensis Lophiotoma cerithiformis Lophiotoma cingulifera Figure 3. t'hylogenetic uee 0f species listed in TaJile 1. Tiiis tree was made using ihe same sequence alignment as that used For the tree in I'igurx.1 1. hut was coiisinietod using ihe l'HYMl. software program,Page 1.38 TH E NAUTILUS, Vol. 121, No. 3 (70%) in the Maximum Likelihood analysis. Given these data, the solution would he to split Lophiotoma into two separate genera, Ijophiatoma and Xctimnrturrls. We feel that at this time the more conservative approach of re­taining the genus Lnphfolomu, mid dividing it into two subgenera, lophiotoma (s.s.) und Xenuroiurm (wilh Lophiatoma (Lophiotoma) acuta and Lophiotoma (Xe- wirolurris) chiguHfura as types, respectively) is prefer­able until a wider range of specics has been sinalyzed. There are u number of Spceie* (presently in Lophiotoma) that are pi'obbituilic to assign (such as Lophiotoma ruth- vcniana (McJvill, 1923)), and we believe that i\ molecular analysis of the«s forms needs to be carried out before wc Fully underhand the relationship between Lophiotoma (s.s.) and Xunuroturri.s. It may well turn out that, when the analysis is completed, tlitr separation between the two branches (b>]>hlotoin/i and Xenuroturris) will be defini­tive; at that point, separating tlie two groups of spceles into different genera will be justified. The major conclusion from this work is that Vnedogemmula should be recognized as a full genus, and is n sister group io the major branch that includes Turrit, Cewiiuclii, «nid Lophloloma (as redefined). ACKNOWLEDGMENTS We thank Sean Christensen for help with experiments Bradford Stevenson for help with the preparation of phy­logenetic trees and Drs. Edgar P. Ilcimer de la Cotera and Estuardo L6pez-Vcr<i for providing the Pobjatiru sample. This work was supported in part by a Program Project grant from the National Institutes of Health, OM4W577. LITERATURE CITED Buuchct, P. i J'. l-ozouet, P. Mucslrati, and V. Ilfrns. 2002. Assessing (be magnitude of specics richness in tropical msiriiic environments: high numbers of molluscs at a New Caledonia site. Biological Journal of the Linnoan Society 75: 421-436. Boudict, P. and J. P. Roeroi. 2005. Classification and nomen­clature) of gastropod families. Malueologia 47; J-397. Bottchet. P.. A. 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