TOWARD COMPREHENSIVENESS: INCREASED MOLECULAR SAMPLING ... · TOWARD COMPREHENSIVENESS: INCREASED MOLECULAR SAMPLING ... Cowries (Gastropoda: Cypraeidae) ... and diversification patterns

TOWARD COMPREHENSIVENESS: INCREASED MOLECULAR SAMPLING

WITHIN CYPRAEIDAE AND ITS PHYLOGENETIC IMPLICATIONS

Christopher P. Meyer

Florida Museum of Natural History, University of Florida,

Gainesville, Florida 32611 USA; [email protected]

ABSTRACT

This paper introduces 73 additional taxa to the existing mitochondrial molecular data-base of 202 taxa for the Cypraeidae and addresses the systematic implications of theirinclusion. Five outgroup members from the Ovulidae are also added. Sequence data areincluded from all previously missing extant named genera (Propustularia, Barycypraea

and Schilderia), completing the overall “generic-level” framework for living cowries. Newlyadded taxa include 47 recognized species, 25 subspecies, and six undescribed taxa. Phy-logenetic results generally are consistent with previous arrangements, with few minor ad-justments. The most significant findings are that: (1) currently recognized Nesiocypraea isbroken into two disparate clades, a deeply rooting Nesiocypraea sensu stricto group andthe more derived Austrasiatica (Lorenz, 1989). (2) Two newly included Barycypraea taxaare sister to Zoila, reaffirming the validity of the subfamilial clade Bernayinae. (3) Theinclusion of a significant number of added Erroneini taxa (N = 24) creates a phylogeneticchallenge because of poor support and recovered relationships inconsistent at first glancewith traditionally recognized affinities. In order to maintain nomenclatural consistency,Erronea is maintained at a generic level, whereas Adusta is dropped to subgeneric statuswithin Erronea. Greater than 90% of currently recognized species are included, and 93%of these are supported by molecular criteria. Moreover, more than 70% of the tested,recognized subspecies are distinct. The phylogeny provides one of the most comprehen-sive, species-level frameworks to date for testing diversification theories in the marinetropics.

Key words: Cypraeidae, molecular systematics, taxon sampling, Cypraea.

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INTRODUCTION

Cowries (Gastropoda: Cypraeidae) are taxo-nomically one of the best known of all mollus-can groups, and have been used frequently toexamine speciation and biogeographic pat-terns in the marine tropics (Schilder, 1965,1969; Foin, 1976; Kay, 1984, 1990; Meyer,2003). A wealth of taxonomic (Schilder &Schilder, 1938, 1971; Schilder, 1939; Lorenz &Hubert, 1993; Groves, 1994; Lorenz, 2002),anatomical (Troschel, 1863; Vayssière, 1923,1927; Riese, 1931; Risbec, 1937; Schilder,1936; Kay, 1957, 1960, 1963, 1985, 1996;Bradner & Kay, 1996; Lorenz, 2000), biogeo-graphic (Schilder, 1965, 1969; Foin, 1976;Burgess, 1985; Liltved, 1989; Lorenz &Hubert, 1993; Lorenz, 2002) and fossil data(Schilder & Schilder, 1971; Kay, 1990, 1996;Groves, 1994) is available for the group; how-ever, what has been lacking is a well-resolved,comprehensive species-level phylogeny.

MALACOLOGIA, 2004, 46(1): 127−156

These phylogenetic hypotheses of relationshipestablish sister pairs at the appropriate taxo-nomic level and provide the framework to testdiversification theories. Meyer (2003) introducedmolecular data for 234 taxa in Cypraeidae andgenerated phylogenetic hypotheses for mostmajor clades as well as sister-group relation-ships for most species. Systematics forCypraeidae were reviewed in light of the resultsand diversification patterns within the tropicswere addressed. The study presented hereinsignificantly increases the comprehensivenessof taxon sampling in the group by introducing 73Cypraeidae and five Ovulidae taxa to the exist-ing molecular dataset and discusses their sys-tematic implications. In addition to broadertaxonomic sampling, this paper presents the re-sults of broader geographic sampling. The ap-pendix lists 147 localities added across thevarious taxa. Five outgroup taxa from six locali-ties are included, and 67 recognized cypraeidspecies or subspecies are added from 75 locali-

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ties. The remaining 66 localities were added tosupposedly known taxa, but revealed six previ-ously unrecognized taxa, some of which maycorrespond to names currently in synonymyupon review of type localities.

MATERIALS AND METHODS

Recognition Criteria: ESU versus OTU

The ultimate goal of this project is to con-struct a comprehensive phylogeny of cypraeidgastropods at the appropriate level for diversi-fication studies. As such, the operational taxo-nomic unit (OTU) chosen for phylogeneticanalyses generally represents an evolution-arily significant unit (ESU) that must fulfillsome minimal criteria established throughgenetic scrutiny. First, mtDNA haplotypes ofsampled individuals must represent a mono-phyletic clade; yet this alone is not sufficient,because any phylogeny has a plethora ofmonophyletic groups, because a clade re-quires only two individuals. Thus, auxiliary cri-teria are required to delineate significant units.Within cowries, these additional criteria are (1)geographic distinction or allopatry, (2) signifi-cant genetic distance from the sister groupsuch that pairwise distance comparisons yielda bimodal distribution, and/or (3) taxonomicrecognition by previous workers. An OTU isincluded in analyses only if at least two ofthese three criteria are met. Most OTUs fulfillall three criteria and are considered evolution-arily significant units (ESUs) (sensu Moritz,1994). These criteria are erected in order todelineate independent evolutionary trajecto-ries, but do not guarantee that the units arereproductively isolated. In a few instances, twoof the three criteria (genetic separation andtaxonomic recognition) are not supported bythe third (exclusive geographic signatures).While the genetic differences (monophyly)between populations indicate some indepen-dent period of evolutionary history betweengeographic regions, it appears that, on occa-sion, haplotypes from outlying regions can mixback into the sister gene pool. The few caseswhere all three criteria are not fulfilled alwaysoccur on the periphery of regions (e.g.,Marquesas, Hawaii) and show asymmetrical,“downstream”, dispersal events (Fig. 1). Ascircumscribed, all ESUs discussed indicateindependent evolutionary histories, but alter-native criteria, such as either nuclear markersor breeding experiments, are needed to verifyreproductive isolation.

FIG. 1. ESU vs. OTU criteria. Phylogram show-ing the relationships among members of thePacific Cribrarula subclade, with bootstrap val-ues for major groups. Four distinct clades areevident, and the names presented on the right:Cribrarula catholicorum, C. gaskoini, C. astaryi,and C. cumingii. Note that single individuals oftwo newly included taxa, C. taitae and C. garciai(white stars), nest within two of the major cladesand show little variation (a single mutation).These two new taxa are introduced as OTUs,because of their distinct morphology and geog-raphy (American Samoa and Easter Island, re-spectively), but are currently not consideredESUs by molecular criteria. All individuals fromthe Marquesas are C. astaryi; however, two in-dividuals of C. cumingii possess haplotypes be-longing to the C. astaryi clade as well (dark stars).While the two haplotype clusters are distinct, thepattern indicates uni-directional exchange of lar-vae downstream from the Marquesas (C. astaryi).Molecular criteria recognize these two clades asESUs with historically limited exchange. (TIK =Tikehau, RR = Rangiroa, HUA = Huahine, all C.astaryi from Marquesas, all C. gaskoini fromHawaii, and all C. catholicorum from SolomonIslands)

Molecular Methods

Most methods follow protocols detailed inMeyer (2003) for all aspects of preservation,extraction, amplification, and sequencing. Tis-sue samples were acquired from a variety of

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sources and locations (listed in theacknowledgements and appendix). Mostsamples were preserved in 95% ethanol. DNAextraction was performed using DNAzol(Chomczynski et al., 1997) using one-half vol-umes and following the manufacturer’s proto-col (Molecular Research Center, Inc.) with theexception that the digestion step was in-creased by an additional 24 or 48 h. PCR wasperformed as described in Meyer (2003). COIprimers were as follow (from Folmer et al.,1994): LCO-1490 (5’−3’) GGT CAA CAA ATCATA AAG ATA TTG G, and HCO-2198 (5’−3’)TAA ACT TCA GGG TGA CCA AAA ATC A.For problematic taxa, these primers were de-generated as follows: dgLCO-1490 (5’−3’)GGT CAA CAA ATC ATA AAG AYA TYG G, anddgHCO-2198 (5’−3’) TAA ACT TCA GGG TGACCA AAR AAY CA. Two internal primers weredesigned for small amplifications of degradedDNA: InCypLCO (5’−3’) CGT YTA AAT AATATA AGY TTY TG, and InCypHCO (5’−3’) CGTATA TTA ATA ATT GTT GTA AT. Palumbi’s(1996) 16Sar and 16Sbr primers were usedfor 16S: 16Sar (5’−3’) CGC CTG TTT ATC AAAAAC AT, and 16Sbr (5’−3’) CCG GTC TGA ACTCAG ATC ACG T. Two internal primers weredesigned for small amplifications of degradedDNA: In16Sar (5’−3’) GGG CTA GTA TGA ATGGTT TGA, and In16Sbr (5’−3’) ATG CTG TTATCC CTA TGG TAA CT. The polymerase chainreaction was carried out in 50 µl volumes, us-ing 1 µl of template. Each reaction included 5µl 10X PCR buffer, 5 µl dNTPs (10mM stock),2 µl of each primer (10µM stock), 3 µl MgCl

2

solution (25 mM stock), 0.2 µl Taq (5 Units/µlstock) and 31.8 µl ddH

2O. Reactions were run

for 35-40 cycles with the following parameters:an initial one min denaturation at 95°C; thencycled at 95°C for 40 sec (denaturation), 40°Cto 44°C (COI) or 50°C to 54°C (16S) for 40 sec(annealing), and 72°C for 60 sec (extension).Successfully amplified products were cleanedfor cycle sequencing using Wizard PCR Preps(Promega). Sequencing also followed Meyer(2003) with all new sequences generated usingABI chemistry and sequencers. Sequences weregenerated from the resulting electrophenogramsusing Sequencher (Gene Codes).

All primer sequences, aligned COI and 16Ssequences and Nexus files are available atthe archived data web pages of the FloridaMuseum of Natural History Malacology Depart-ment (http://www.flmnh.ufl.edu/malacology/archdata/Meyer2004), and new sequences aredeposited in Genbank under accession num-bers AY534351 through AY534503.

Phylogenetic Analyses

The 297 operational taxonomic units (OTUs)presented in this paper were selected from anextensive database comprised of over 2,000sequenced individuals. In general, taxa areincluded if they exhibit distinctive geographicand/or genetic signatures. In most instances,new OTUs are recognized in the literature aseither species (N = 47) or subspecies (N =25). This paper introduces six previously un-recognized taxa.

The increasing size of this dataset presentscomputational and heuristic challenges forphylogenetic analyses. Two weighted trans-version bias parsimony searches (3:1 and 5:1)were performed on the complete dataset us-ing PAUP* (Swofford, 1998). At first, 250 ran-dom-addition replicate searches wereperformed, but with a tree limit of ten imposedto minimize search time on suboptimal islands.After 250 replicates, the most parsimonioustopologies were used as starting trees for ex-haustive searches without tree limits. Thisstrategy was employed for both weightedanalyses, and the most parsimonious topolo-gies were pooled and evaluated using likeli-hood criteria. ModelTest v. 3.06 (Posada &Crandall, 1998) was used to select the mostappropriate model for likelihood parameters.The most likely weighted parsimonious treeswere then compared using consensus meth-ods.

A two-tiered, compartmentalized strategywas adopted that followed Meyer (2003) forlevels of topological support. The strict con-sensus topology derived from the most likelyoverall analyses was divided into foursubequal components called basal, mid1,mid2, and derived. Because the basal, mid1and mid2 cohorts are necessarily paraphyleticgroups that include the common ancestor andsome, but not all, of its descendants, repre-sentative derived clades were included in theparaphyletic analyses. In this way multiplederived member clades overlapped betweenmore basal and derived analyses, and theoverall topology could be “scaffolded” togetherby linking clades shared in both basal andderived compartments.

Within each of the four subanalyses, parsi-mony searches were performed using a 5:1transversion bias. Both bootstrap (Felsenstein,1985) analyses (1,000 replicates) and decay(Bremer, 1994) analyses (TreeRot v2;Sorenson, 1999) were performed to establishlevels of support. Results from Bayesian meth-

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ods (Mr. Bayes v3.04b) are not reported in thispaper, but were generated for the four sub-groups and compared to the combined parsi-mony/likelihood methods utilized in PAUP*.Overwhelmingly, they were consistent with theresults presented here, but on few occasionsdiffered in hypotheses of relationship. Thescaffolded parsimony global topologies werecompared to the scaffolded Bayesian topol-ogy using likelihood criteria in PAUP*. Thecombined topology derived from the compart-mentalized Bayesian subsets was less likelythan the overall topologies found using thecombined parsimony/likelihood criteria. It ap-pears that Bayesian results depended on taxonsampling and outgroup inclusion. While thisfinding may be of interest to the general sys-tematic community, it is not a point specificallyaddressed in this paper.

RESULTS

The final culled dataset contained 297 OTUsand 1,107 characters, 493 base pairs from 16Sand 614 bases from COI. For 16S, alignmentfollowed those presented in Meyer (2003)based on secondary structure. Weighted par-simony searches resulted in 512 equally mostparsimonious trees (MPTs) for 3:1 Ti:Tv and 480trees for 5:1 searches. Derived portions of thecomprehensive topology were consistent. Thus,all named clades (subfamilies, tribes and gen-era) presented in Figure 2 are found in all to-pologies, except one mentioned below.However, the topologies recovered from alter-nate weightings differed in five deeper regions,all of which are poorly supported regardless ofmethodology. First, 5:1 topologies placed theclade consisting of Propustularia/Nesiocypraea/Ipsa basal as sister to all other cowries. In 3:1topologies this clade moves up one node andis sister to Erosariinae. Second, the pustuloseclade consisting of Nucleolaria/Cryptocypraea/Staphylaea is monophyletic in 5:1 trees, whilein 3:1 topologies these genera are a basalparaphyletic grade leading to the clade includ-ing Monetaria/Perisserosa/Erosaria. Third, in5:1 topologies Perisserosa is sister to Erosaria,whereas in 3:1 trees, Perisserosa is sister toMonetaria. Fourth, the arrangement of majorgroups along the backbone from Umbiliini toCypraeovulinae conflicts. Results from 5:1searches are shown in Figure 2, whereas in3:1 topologies, Notocypraea and Cypraeovula(Cypraeovulinae) are a basal sister grade lead-ing to more derived member groups. Finally,the basal arrangement within Erroneini is dif-

ferent. In 3:1 topologies Purpuradusta is morebasal, while in 5:1 trees, Erronea is more basal.

When alternative topologies were evaluatedusing ModelTest, the GTR+I+G model was se-lected as the best-fit model. When both the 3:1MPTs and 5:1 MPTs were evaluated using theselected likelihood criteria [lset base =(0.315128 0.136452 0.111915), Nst = 6,Rmat = (0.99559 41.36057 1.0461 1.6893522.78834), rates = gamma, shape = 0.562423,Pinvar = 0.48426], the 5:1 subset was signifi-cantly more likely (ANOVA: p < 0.001, average–ln likelihood = 49513.8). Therefore, resultsfrom the 5:1 searches are presented herein.

The overall relationships among major sub-groups recovered in the 5:1 MPTs are moreconsistent with both morphological and fossilevidence in addition to being more likely basedon molecular data. In particular, a monophyl-etic pustulose clade is more parsimonious forconchological and anatomical features, be-cause it is more likely that a bumpy shell wasderived a single time, rather than being derivedeither twice independently, or derived once thenlost. Also, the basal, paraphyletic status ofNotocypraea and Cypraeovula within the 3:1topologies is inconsistent with the fossil recordfor both groups relative to more derived mem-bers of the 3:1 MPTs (i.e., Umbilia, Barycypraea,and Zoila), which appear earlier in the recordand root more deeply in the 5:1 topologies. Also,the sister-group relationship of the two generais more consistent with paleobiogeography (thebreakup of Gondwanaland) and recognizedaffinities based on both conchological and de-velopmental criteria. The other major discrep-ancies between the 3:1 and 5:1 MPTs (mostbasal cowries, Perisserosa affinities, and posi-tion of Purpuradusta) are more ambiguousbased on alternate criteria (morphological orpaleontological).

Suprageneric Relationships (Fig. 2)

Overall, suprageneric results were consistentwith previous systematic findings (Meyer,2003), with two exceptions. First, Ipsa falls out-side Erosariinae and is no longer sister toErosariini, but instead is allied with newly in-cluded Propustularia and Nesiocypraea sensustricto. New sequence data from Nesiocypraeateramachii neocaledonica did not result in anaffinity with other recognized “Nesiocypraea”species (N. hirasei, N. sakurai and N. langfordi).Instead, Nesiocypraea teramachii roots moredeeply in the phylogeny as a distant sister toIpsa childreni, within a clade that includes bothIpsa and Propustularia. Thus, the inclusion of


FIG. 2. Strict suprageneric consensus topology of 480 most parsimonious trees derived from a5:1 Ti:Tv weighted search strategy of all 297 OTUs. Subfamilies are indicated with arrows andtribes are listed to the right. The four compartments for further subanalyses are bracketed to theright. The four newly added genera are capitalized and bolded. 1Lyncina includes the subcladesCallistocypraea, Miolyncina and Lyncina as reported in Meyer (2003). 2Austrasiatica replacesthe prior use of Nesiocypraea for the same clade. 3Erronea now includes Adusta, formerly rec-ognized as the sister taxon.

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two new ancient lineages (Propustularia andNesiocypraea) affects the relative position ofIpsa. Moreover, the finding that Nesiocypraeateramachii is not related to other previously rec-ognized Nesiocypraea, compels me to recog-nize the clade Austrasiatica proposed by Lorenz(1989) at the generic level for the group includ-ing Austrasiatica hirasei, A. sakurai, and A.langfordi. There are some conchological andanatomical features that support this separa-tion. The left posterior terminal ridge inNesiocypraea is more produced and separatefrom the body of the shell, whereas inAustrasiatica, the ridge is continuous with thebody. Lorenz (pers. comm.) also states that (1)Nesiocypraea lacks a distinct embryonic band-ing, having instead only a darker middorsalzone, (2) Nesiocypraea have a proportionallylarger spire, and (3) the darker pattern of theshell is absent in juvenile Austrasiatica, onlygained after the deflection of the labral margin;whereas, the darker pattern can be part of ju-venile Nesiocypraea shells. Additionally, therachidian tooth of Nesiocypraea lacks theprominent paired basal denticles present in thethree Austrasiatica taxa, and the tooth shapeis less elongated and squared, whereas therachidian in Austrasiatica narrows toward thecusps (Bradner & Kay, 1996). The fact thatAustrasiatica was erected to differentiate thethree species (albeit incorrectly aligned withSchilderia) is also an indication that the two lin-eages possess independent histories. The deepposition of Propustularia within the cowrie phy-logeny is not surprising because it is one of theoldest of extant taxa, extending back to theLower Eocene (Kay, 1996).

The second suprageneric difference concernsthe relative position of Zoila in the overall phy-logeny and is caused by the inclusion of se-quence data for two taxa from the ancientlineage Barycypraea. These new data indicatethat Barycypraea teulerei and Barycypraeafultoni are sister taxa, and they are sister toZoila. This Barycypraea/Zoila clade is recog-nized as the extant members of the subfamilyBernayinae, a group that includes many extinctfossil members and extends back into the Me-sozoic (Kay, 1996). These new data changethe relative position of Zoila to Cypraeinae(Meyer, 2003); however, the topology in thisregion of the phylogeny is poorly supported.

The final suprageneric addition to the molecu-lar database is the inclusion of sequence datafrom Schilderia achatidea, the single, living rep-resentative from an older, more diverse genusof European affinities. Previously, theparaphyletic arrangement of the genera

Pseudozonaria and Zonaria was a surprisingresult (Meyer, 2003). These new data forSchilderia place the genus as sister to Zonariato the exclusion of Pseudozonaria (andNeobernaya), and phylogenetic results main-tain their independent, paraphyletic status.These finding are more consistent with geo-graphic affinities than recognized taxonomicaffinities (Pseudozonaria is often considered asubgenus of Zonaria), as both Neobernaya andPseudozonaria are currently restricted to theeastern Pacific whereas Schilderia and Zonariaare restricted to the western Atlantic.

Basal Compartment (Fig. 3)

Five Ovulidae taxa are added in these analy-ses: Pseudocypraea exquisita, Volva volva,Primovula concinna, Dentiovula takeoi, andProsimnia semperi. Within Ovulidae, only a fewmajor clades are well supported and may bethe results of poor taxon sampling. First, theclade Eocypraeinae appears well supportedand includes Pedicularia, Jenneria andPseudocypraea. Eocypraeinae is sister to astrongly supported clade (Ovulinae) that in-cludes the remaining Ovulidae. Within theOvulinae, two subgroups are well supportedand represent the major clades Volvini andOvulini. Of the added Ovulidae, Volva falls intoVolvini, but Prosimnia unexpectedly falls intoOvulini as do Primovula and Dentiovula. Theseresults are generally consistent with Cate’s(1974) arrangement of higher-level relation-ships within the Ovulidae. Cyphoma gibbosumfalls basal to these two sisters in the strict con-sensus topology; however, its position is poorlysupported, and it is expected to move withinthe Volvini with the inclusion of more taxa.Monophyly of Ovulidae is not addressed hereinand would require the inclusion of more distantrepresentatives from Lamellaridae, Triviidaeand Eratoidae.

The Cypraeidae basal group includes thegenera Propustularia, Nesiocypraea, Ipsa,Cryptocypraea, Nucleolaria, Staphylaea,Monetaria, Perisserosa, and Erosaria.Propustularia, Nesiocypraea, and Ipsa form aclade that roots deeply within the phylogeny andis sister to all other cowries. Each of the threegenera is represented by only a single taxon,and only Nesiocypraea contains additional rec-ognized species missing from the dataset(Nesiocypraea midwayensis, N. lisetae and N.aenigma). While sharing a most recent com-mon ancestor, the three genera are highly di-vergent from each other, representingsignificant periods of independent history. Two


FIG. 3. Basal Compartment cladogram and phylogram. Bootstrap values are presented abovebranches in the cladogram and rescaled decay values below. Bolded taxa are new additions to thedata set. Their identity number shown in parentheses follows the listing in the Appendix. Generic orsuprageneric groupings are indicated to the right of the cladogram. OTUs with an asterisk (*) are notESUs based on molecular criteria. Phylogram to the right is based on likelihood distances using aGTR+I+G model of sequence evolution. Note that the scaling for branch lengths changes betweenOvulidae and Cypraeidae.

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are known exclusively from the Indo-Pacific(Nesiocypraea and Ipsa) and one(Propustularia) from the western Atlantic, buthas a fossil record from North America, theCaribbean, and Europe (Kay, 1996). The splitsamong these ancient groups are among theearliest of all extant species and may have oc-curred in the Mesozoic. While reasonably sup-ported as a clade, this basal group is notstrongly supported as the most basal sister, andin other analyses (3:1) moves up to becomesister of the remaining basal taxa (Erosariinae).

The final six genera from the basal compart-ment form the strongly supported cladeErosariinae and is the sister group to all remain-ing extant species. Membership and relation-ships within the Erosariinae are consistent withprevious findings (Meyer, 2003). Five taxa fromErosaria are added: Erosaria marginalis, E.citrina, E. helvola cf. callista, E. macandrewi,and E. englerti. Ten independent lineages arestrongly supported (bootstraps > 90/decays >6) within Erosaria, but interrelationships amongthem are not (< 50/< 4). Erosaria marginalisand E. citrina, both from the western IndianOcean, are strongly supported as sister taxa.This clade is poorly supported as sister to theE. helvola complex. Within Erosaria helvola,three ESUs are identifiable: E. helvolahawaiiensis from Hawaii, E. helvola cf. callistafrom the Marquesas, and E. helvola helvolafrom the remainder of the IndoPacific. Thenewly included ESU, E. helvola cf. callista, mayneed a new name, because the type locality ofE. helvola callista is Tahiti (Shaw, 1909), notthe Marquesas. These five taxa are sister tothe remaining Erosaria; however, the basal po-sition is poorly supported. Erosaria turdus is amonotypic, deeply divergent lineage. Newlyadded Erosaria irrorata, a species restricted tothe oceanic islands of the Pacific, is poorly sup-ported as sister to a strongly supported clade(97/12) including E. albuginosa and E. poraria.These three taxa are sister to a well-supportedlineage (92/6) of eight taxa that I tentatively rec-ognize as Paulonaria at the subgeneric level.New sequence data from Erosaria macandrewi,a Red Sea taxon, closely ally that species withE. beckii. These two species are sister to theremaining Paulonaria taxa. The final additionaltaxon within Paulonaria is Erosaria englerti, aspecies endemic to Easter Island and Sala yGomez. Erosaria englerti shares a more recentcommon ancestor with the remaining fivePaulonaria taxa. All other relationships withinErosaria are the same as those presented inMeyer (2003) and are indicated in Figure 3.Newly added haplotypes from E. lamarckii

lamarckii populations of the western IndianOcean exhibit a recent divergence from thepreviously recorded E. lamarckii cf. redimita ofthe Andaman Sea. One final finding from addi-tional Erosaria sequence data is that haplotypesfrom Erosaria miliaris and E. eburnea individu-als interfinger, indicating that either the diver-gence between these two taxa is very recentand lineage sorting has not occurred, or thatthese two taxa represent a cline across thewestern Pacific from a colored dorsum in thewest to white shells in the east.

Mid1 Compartment (Fig. 4)

The second paraphyletic compartment con-tains mostly large-shelled taxa from the follow-ing tribes: Umbiliini, Cypraeini, Mauritiini, Luriini,Austrocypraeini, and the genus Pustularia. Allsix clades are well supported (> 70/> 5) exceptfor Austrocypraeini. As in Meyer (2003), inter-relationships among these major supragenericclades are resolved in the consensus, but poorlysupported. Austrocypraeini and Luriini are sis-ters and recognized as the subfamily Luriinae.Barycypraea and Zoila are sisters and recog-nized as the subfamily Bernayinae. Cypraeiniand Mauritiini are sisters and recognized as thesubfamily Cypraeinae. In the current topology,Pustularia and all remaining cowries share amore recent common ancestor. This large cladeis sister to Luriinae, which in turn is sister toBernayinae, and this inclusive clade is sister toCypraeinae. As in Meyer (2003), Umbiliini is sis-ter to all remaining mid1, mid2 and derived taxa.

Within the mid1 compartment, 13 taxa areadded to the sequence database. The first addi-tion falls within the genus Umbilia and is tenta-tively recognized as Umbilia cf. petilirostris. Asingle divergent sequence was generated fromtissue samples collected from the deep watersin the Capricorn Channel off Queensland, Aus-tralia. Seven sequenced individuals were com-pletely identical, while an eighth sample from asubadult shell was significantly divergent. Thissingle sample may represent the newly de-scribed Umbilia petilirostris Darragh, 2002; how-ever, authors disagree on its taxonomic status(Wilson & Clarkson, in press). Until more com-prehensive sampling is done in the region, Ipresent the divergent sequence as a differentESU, which does not preclude it from beinglumped within U. capricornica at a later datewith more exhaustive sampling. The relation-ships within Umbilia remain as in previousanalyses (Meyer, 2003).

The second taxon added to mid1 is Lepori-cypraea mappa aliwalensis from Natal, South


FIG. 4. Mid 1 Compartment cladogram and phylogram. All other information as in Fig. 3.

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Africa, and falls as sister to Leporicypraeamappa rosea. Lorenz (2002) has recently re-vised the taxonomy of the mappa group in lightof molecular findings. Importantly, the names Iassociated previously with ESUs havechanged, and those changes are reflected inthe Appendix and also discussed herein. Thetaxon I previously recognized as Leporicypraeamappa viridis from SE Polynesia is now recog-nized as Leporicypraea admirabilis. The taxonI previously recognized as Leporicypraeamappa panerythra from the non-continentalportions of the western Pacific is now recog-nized as Leporicypraea mappa viridis. The othertaxon names remain the same. Sequences ofL. mappa “rewa” from Pacific localities (Fiji,Vanuatu, Palau, and South China Sea)interfinger with haplotypes of L. mappageographica individuals from Indian Ocean lo-calities (NW Australia, Phuket, Seychelles, andZanzibar). Therefore, I recognize only a singletaxon, L. mappa geographica, for this clade.Because of its conchological distinctivenessand sympatry with conspecifics, Lorenz (2002)elevated L. mappa geographica to specific sta-tus with Indian and Pacific subspecies. Basedon the genetic difference between mappa-com-plex conspecifics and geographic overlap, spe-cific status is certainly acceptable. However, theremaining L. mappa subspecies are para-phyletic. The phylogeny Lorenz (2002: 27) pre-sents is correct and reflects this arrangement.Certainly, other recognized cowrie species arederived from paraphyletic parent species (e.g.,Eclogavena coxeni and others; see Meyer,2003: table 4, and cases herein), and L.geographica would have to be added to thislist. These results suggest a third species sis-ter to L. geographica should be recognized thatwould include both L. mappa viridis and L.mappa admirabilis. L. mappa geographica in-dividuals have been found sympatrically withboth L. mappa mappa and L. mappa viridis in-dividuals in the Pacific Ocean. However, as yet,L. mappa mappa and L. mappa viridis haplo-types have not been found together.

One new undescribed taxon is added toMauritia. Haplotypes of M. arabica individualsfrom American Samoa cluster independentlyfrom haplotypes of M. arabica individuals fromother Pacific localities. Shells from Samoan in-dividuals tend to be smaller, more heavily mar-gined and more circular than individuals fromother Pacific localities. Results from increasedsampling in both M. depressa depressa (N =10) and M. depressa dispersa (N = 10) main-tain their independent, reciprocally monophyl-etic status, albeit recently diverged. As in

previous findings, the interrelationships amongmajor lineages in Mauritia are poorly supported.Consensus methods and poor support resultin two polytomies (Fig. 4). Further genetic datawill be needed to address this region of the phy-logeny as all extant taxa have been sampled.

New sequence data from Barycypraeateulerei and B. fultoni place them as sister taxaand align them with the genus Zoila to form thegroup Bernayinae. Sequence data presentedfor Barycypraea fultoni are of B. fultoni amorimifrom Mozambique. The Australian Zoilamarginata complex is split into two ESUs asincreased sampling indicates fixed moleculardifferences between populations separated bythe Southwest Cape region between capesNaturaliste and Leeuwin. Further sampling di-rectly within this region may uncover interme-diate haplotypes that would link the two ESUsand suggest a cline instead of two independentlineages. Such a finding is the case in the Zoilafriendii complex. However, as none have beendiscovered yet, I present the data as two tenta-tive ESUs: Zoila marginata marginata to thesouth and Z. marginata ketyana to the west.Other described Z. marginata taxa (Lorenz,2001; 2002) within each ESU interfinger, anddo not fulfill molecular criteria for recognition.Sequence data from Zoila mariellae are the fi-nal addition to the Bernayinae clade. While theexact provenance of the animal sequenced isunknown, it is likely from the northwestern shelfof Australia. Molecular results place Z. mariellaeas a distinct sister to Z. decipiens, also fromthe northwestern shelf, as expected.

Following along the phylogeny, the cladeLuriinae comes next. Talparia and Luria arestrongly supported as the clade Luriini. A smallfragment from 16S was amplified from a de-graded Talparia exusta specimen, and as ex-pected, the taxon is sister to the morewidespread Talparia talpa. Surprisingly, se-quence divergence between the two speciesappears to be relative small, indicating a morerecent divergence than expected. Better-pre-served material from T. exusta is needed be-fore these relative results can be confidentlyassessed. The inclusion of four new taxa to theAustrocypraeini (Arestoides argus contrastriata,Lyncina broderipii, L. ventriculus from the In-dian Ocean, and L. kuroharai) does not help inresolving interrelationships among membertaxa. Arestoides argus is broken into a Pacificclade, A. argus argus, and a western IndianOcean clade, A. argus contrastriata, based onadditional sequence data from the IndianOcean. Lyncina broderipii appears as sister toL. nivosa within the Callistocypraea clade, as


predicted in Meyer (2003). A single sampledindividual of L. ventriculus from Christmas Is-land in the Indian Ocean falls significantly out-side the haplotype cluster of individuals (N = 6)from various regions of the Pacific basin.Lyncina ventriculus is an oceanic taxon, andbecause of the geographic gap between sitesacross continental Southeast Asia, I choose topresent the Christmas Island form as new,undescribed, distinct ESU. Further sampling ofindividuals from Christmas Island may changethis interpretation, but they are currently lack-ing. A single sample of Lyncina kuroharai wassequenced and the results place it closely re-lated to L. sulcidentata, an endemic Hawaiiantaxon. The shallow split between these two taxaindicates a relatively recent common ancestor.Faunal ties have been documented in othercowrie species between Hawaii and Japan,most notably in Luria isabella, and the closeaffinities between L. kuroharai and L.sulcidentata represent another example of thisbiogeographic link.

The final two ESUs added within the mid1compartment are members of the genusPustularia, and more specifically are recognizedsubspecies of Pustularia bistrinotata. A singleP. bistrinotata keelingensis individual was se-quenced, is distinct, and appears as sister tothe remaining P. bistrinotata complex. Further-more, P. bistrinotata sublaevis individuals (N =5) from southeast Polynesia (Tuamotu andSocieties) cluster together, forming a third ESUwithin P. bistrinotata.

Mid2 Compartment (Fig. 5)

The third phylogenetic compartment, mid2,contains members from the generaNeobernaya, Pseudozonaria, Schilderia,Zonaria, the subfamily Cypraeovulinae, and thetribe Bistolidini of the subfamily Erroneinae.Interrelationships among member clades areconsistent with previous findings (Meyer, 2003).Neobernaya and Pseudozonaria are sisters,and that clade is sister to the remaining cow-ries. The inclusion of sequence data from thegenus Schilderia (S. achatidea), place thegroup as sister to Zonaria, and together thisclade shares a more recent ancestor with theremaining taxa. The subfamily Cypraeovulinaeincludes both the South African Cypraeovulaand South Australian Notocypraea and is sis-ter to the western IndoPacific Erroneinae, whichis composed of two tribes: Bistolidini andErroneini.

Within the mid2 compartment, 25 taxa areadded to the existing sequence database; at

least one ESU is added within each genus ex-cept the monotypic Neobernaya. Pseudo-zonaria nigropunctata, a Galapagos endemic,falls into the eastern Pacific clade as a diver-gent sister to P. arabicula, although not stronglysupported. The position of Schilderia achatideahas been mentioned previously as sister toZonaria, now found exclusively in the easternAtlantic. Two taxa are added from Zonaria.Zonaria picta from the Cape Verde Islands fallsnear the base of Zonaria, and its relationshipwith other Zonarid taxa is ambiguous, resultingin a polytomy at the base of the group. Alterna-tive phylogenetic reconstructions at the baseof the group show small internodes, indicativeof a short radiative burst, with little divergencesince. New sequence data from Pseudozonariaangelicae are extremely similar to haplotypesfrom P. pyrum (both P. pyrum angolensis andP. pyrum senegalensis). I include P. angelicaeas a taxon in the phylogeny, but prefer to con-sider it at most a subspecies until further se-quence data are available within the P. pyrumcomplex, as I have reservations concerning di-vergences along the mostly continuous WestAfrican/Mediterranean coastline.

Sequence data from six additional taxa areincluded within Cypraeovulinae, two fromNotocypraea and four from Cypraeovula. InNotocypraea, I tentatively recognize two ESUswithin Notocypraea angustata, with a phyloge-netic break somewhere between Port Lincolnand Port Macdonnel, South Australia. Two di-vergent haplotype clusters exist without inter-mediate states. Again, further data may changethis interpretation, but at present I chose to rep-resent these as different ESUs indicating dis-tinct evolutionary trajectories. Sequence datafrom a single specimen of Notocypraeahartsmithi, a rare species from southeasternAustralia, indicate that the species is sister toall remaining Notocypraea taxa. WithinCypraeovula, four taxa are added, but their in-clusion does not change previous interpreta-tions that the group is composed ofpredominately four divergent lineages with mi-nor differences within each. New sequence datafrom both Cypraeovula fuscorubra and C.fuscodentata closely align these taxa with C.capensis. New sequence data from C. mikehartiand C. algoensis closely align those taxa withC. edentula and C. alfredensis. Noting the shal-low divergences among recognized species inFigure 5, I am doubtful that many of the de-scribed subspecies within Cypraeovula (sum-marized in Lorenz, 2002) will fulfil l mymolecular criteria for ESU status. As somespecies are differentiated currently by only a

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FIG. 5. Mid 2 Compartment cladogram and phylogram. All other information as in Fig. 3.


single mutation (e.g., Cypraeovula mikeharti/C. algoensis or C. castanea/C. iutsui), theresimply is not enough room for differences tohave accumulated between taxa. This is notto say that described entities are not indepen-dent. Indeed, because Cypraeovula taxa aredirect developers with limited dispersal andgene flow, regional differences are expectedon small geographic scales, much like theSouth Australian endemic clades Umbilia,Zoila, and Notocypraea. However, based on thegenetic similarity among sampled memberCypraeovula, much of this variation has to bevery recently derived. This pattern is borne outin the South Australian direct developers thathave been more extensively sampled.

The tribe Bistolidini within Erroneinae is com-posed of members from five genera:Palmadusta, Bistolida, Ovatipsa, Talostolidaand Cribrarula. As in Meyer (2003), the basalroot of Bistolidini is poorly resolved. Overallanalyses place either Palmadusta as sister tothe other four genera or Palmadusta andBistolida as a clade, sister to the remainingthree. Compartmentalized analyses placePalmadusta at the base, although poorly sup-ported. The addition of 15 ESUs did not help inresolving this issue. Only one taxon is addedto the Palmadusta clade, but it alters the sub-specific designations previously ascribed(Meyer, 2003). New haplotypes from AndamanSea P. clandestina individuals form a distinctmonophyletic clade. This new ESU is sister tothe western Indian Ocean P. clandestinapasserina, and the two of them are sister to thePacific P. clandestina clade and the Japaneseendemic P. artuffeli. Based on a review of P.clandestina subspecies and type localities, thePacific clade that I had formerly (Meyer, 2003)recognized as P. clandestina clandestina shouldbe P. clandestina candida, and the new P.clandestina clade from the Andaman Sea nowbears the name P. clandestina clandestina. Ialso reviewed the subspecies and type locali-ties for the three P. asellus ESUs previouslyunnamed (Meyer, 2003). Based on increasedsampling and conchological comparisons, I ten-tatively ascribe the following subspecific des-ignations for the three clades: P. asellus asellusfor the western Indian Ocean clade, P. asellusvespacea for the Seychelles to western Pacificclade, and P. asellus bitaeniata for theMelanesian and Pacific clade (Fig. 5, Appen-dix). Unfortunately, the addition of P. clandestinaclandestina does not help in resolving the basalnodes of Palmadusta. As shown in Figure 5,the base of Palmadusta is poorly resolved andsister group assignments are ambiguous. A few

lineages remain strongly supported (P. asellus,P. clandestina/diluculum, P. ziczac and P.contaminata), but confident hypotheses of otherinterrelationships require further data.

Three taxa are added to Bistolida: B. stolidadiagues, B. owenii and an undescribed, dis-tinct eastern Indian Ocean clade of B. ursellus.Individuals of B. stolida diagues from theSeychelles fall as sister to B. stolidarubiginosa. Bistolida owenii, a western IndianOcean taxon, is sister to the Red Sea endemicB. erythraeensis. A new Bistolida ursellus se-quence from the Andaman Sea is poorly sup-ported as sister to the remaining B. ursellustaxon from the Pacific basin. Its placement isequally parsimonious as either sister to B.ursellus (Pacific) or forming a B. ursellus gradeleading to the B. kieneri lineage. The topologyof the two B. ursellus taxa as sisters is morelikely and consistent with morphology.

One taxon is added to Ovatipsa and two taxato Talostolida. Within Ovatipsa, the subspe-cies O. chinensis amiges from the Pacific ba-sin and Western Australia is distinct from O.chinensis chinensis from the Philippines west-ward through the Indian Ocean to the eastcoast of Africa. Various other O. chinensissubspecies have been described within theIndian Ocean (e.g., Lorenz & Hubert, 1993),and preliminary data indicate that these IndianOcean subspecies may represent very recentdivergences within what I am currently recog-nizing as O. chinensis chinensis. However,until more individuals are sampled, I maintainthem all under the taxon Ovatipsa chinensischinensis. Within Talostolida, two taxa areadded that appear as sisters to each other: T.subteres from southeastern Polynesia and T.latior from Hawaii. These two taxa are sisterto Talostolida pellucens. All four taxa currentlyincluded within Talostolida are deeply divergentindependent ESUs. A single haplotype ofTalostolida teres “alveolus” (sensu Lorenz,2002) is completely identical to haplotypes ofT. teres teres individuals from both the SocietyIslands and the Tuamotu. Moreover, T. teresindividuals from SE Polynesia have been de-scribed by Lorenz (2002) as a distinct subspe-cies T. teres “janae”; however sampledindividuals of T. teres from SE Polynesiainterfinger with individuals sampled from theWestern Pacific (Papua New Guinea andGuam). Therefore, the data do not support T.teres “janae” as a valid taxon, based on mycriteria. All Marquesan individuals sequencedpossess T. pellucens haplotypes, whereas allT. teres-like individuals from the remainder ofSE Polynesia possess T. teres haplotypes.

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The Cribrarula clade includes eight additionaltaxa, making it the most diverse genus withinBistolidini. Two taxa, Cribrarula taitae fromAmerican Samoa and C. garciai from EasterIsland, are added to the deeply divergent Pa-cific subclade. Both taxa are recently divergentmembers from their respective sister taxon.Cribrarula taitae appears as a closely relatedsister to C. catholicorum, and C. garciai isclosely related to C. cumingii. Only a single in-dividual from each of the two taxa was includedin these analyses, and the results would bebetter addressed with multiple samples. Twomembers are added to the Western IndianOcean subclade: Cribrarula pellisserpentis andC. esontropia francescoi, both from Madagas-car. Cribrarula esontropia francescoi is a closelyrelated sister to C. esontropia esontropia, whichincludes C. esontropia cribellum (Meyer, 2003).Cribrarula pellisserpentis is a deeply divergentmember within the western Indian Oceansubclade and is sister to the other three ESUs.Four taxa are added to the remaining Cribrarulamember clade. A single individual of C. cribrariafrom Masirah, Oman, appears significantly di-vergent from population samples of the previ-ously unnamed C. cribraria ESU from theAndaman Sea. Conchologically, this individualapproximates the western Indian Ocean taxonC. cribraria abaliena and is tentatively recog-nized as such. A single individual of C. cribrariaaustraliensis from Western Australia falls withinthe Andaman C. cribraria cluster; therefore, Itentatively adopt the name C. cribraria cf.“australiensis” for a taxon that extends from theAndaman Sea southward to Western Austra-lia. More exhaustive sampling is required toconfirm these geographic patterns. A single in-dividual of C. exmouthensis magnifica fromBroome is significantly different from samplesof C. exmouthensis exmouthensis from theExmouth Gulf region, therefore validating thestatus of that taxon. Additional samples of C.cribraria rottnestensis (N = 3) further validatethe taxon’s uniqueness. Eight individuals of C.melwardi from northeastern Australia all sharea common ancestor and are reciprocally mono-phyletic with respect to the remaining C.cribraria individuals. Moreover, a single C.cribraria cribraria individual from the same reef(Lamont Reef in the Bunker Group) clusters asexpected with other Pacific C. cribraria cribrariaindividuals. The final taxon included is C.cribraria abrolhensis (N = 3), and haplotypesare shallowly divergent but reciprocally mono-phyletic with respect to samples of C. cribrariacribraria (N = 30) from predominately westernPacific localities (Appendix). More thorough

analyses and discussion of this fascinating,species-rich group is in preparation (Meyer etal., in prep.).

Derived Compartment (Fig. 6)

The final compartment analyzed is the derivedmonophyletic clade recognized as the tribeErroneini. This clade includes the following ninegenera: Austrasiatica, Palmulacypraea,Erronea, Purpuradusta, Contradusta, Nota-dusta, Eclogavena, Melicerona and Blasicrura.Many (25) taxa are added within the tribe, andphylogenetic analyses result in some surpris-ing affinities. For the most part, major generaare well supported, but their interrelationshipsare not. Three taxa currently ascribed toAustrasiatica were included in previous analy-ses (Meyer, 2003); however, they were consid-ered as representatives of the genusNesiocypraea. As discussed earlier, the find-ing that Nesiocypraea teramachii is distantlyrelated raises the subgenus Austrasiatica togeneric status for the clade that includesAustrasiatica langfordi, A. hirasei and A.sakurai. As in Meyer (2003), Austrasiatica issister to all other Erroneini taxa, followed byPamulacypraea as sister to the remainder. Aspredicted in Meyer (2003), the newly addedPamulacypraea musumea falls as sister to P.katsuae. Even with the addition of 24 taxa (a67% increase), the topology among the rest ofthe major Erroneini lineages is ambiguous. Sixadded “Erronea” species form a basal gradeleading to the Adusta/Erronea split previouslyrecognized in Meyer (2003). I take a conserva-tive approach and redefine Erronea to includeall these taxa and subsume Adusta to a well-supported subclade within the group, as thenew data demonstrate that Adusta and Erronea(including the more recent additions) are notequivalent (sisters). If Adusta were to be main-tained at equivalent generic status, Erroneawould represent a paraphyletic group.Purpuradusta, Eclogavena, Melicerona andBlasicrura are all well-supported monophyleticlineages. As in Meyer (2003), Notadusta is wellsupported only if restricted to members of theNotadusta punctata complex. However, be-cause Notadusta martini is often considered amember of Notadusta, I include it withinNotadusta here, although poorly supported. Ina similarly conservative manner, I include twoof the added taxa within Contradusta, althoughagain poorly supported. Support for relation-ships among these seven genera is poor andis likely because of the short internode lengthbetween divergent lineages.


FIG. 6. Derived Compartment cladogram and phylogram. All other information as in Fig. 3.

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Twelve additional taxa are added to Erronea.Six of the additions are traditionally recognizedas distinct species, four have been recognizedas subspecies, and two are newly discovered,but may have names associated with them thathave been placed into synonymy. Of the newspecies, three form a relatively well-supportedclade: Erronea rabaulensis shares a more re-cent common ancestor with E. fernandoi (80/1),and those two are sister to E. vredenburgi (84/3). The three additional Erronea species all nestdeeply within the clade, and their relationshipsare not well supported. Erronea pallida appearsas sister to the clade of the previously describedthree species and Adusta. Erronea pyriformisis relatively well supported (81/6) as the sisterto the clade previously recognized as Erronea(Meyer, 2003). Finally, Erronea xanthodon fallsat the base of Erronea and is sister to all otherErronea taxa. Within the crown Erroneasubclade, six taxa are added that are all tradi-tionally recognized at the subspecific level. In-dividuals of Erronea cylindrica lenella (N = 8,all from New Caledonia) form a monophyleticgroup strongly supported (91/6) as sister to theclade including the remaining E. cylindrica in-dividuals plus two subspecies of E. ovum.These results imply that E. cylindrica at thespecific level is a paraphyletic taxon. Newlyadded individuals of Erronea ovum ovum fromboth Singapore and the Philippines (N = 15)form a monophyletic group sister to E. ovumpalauensis (N = 7). The four remaining, newlyadded taxa are all members of the Erroneacaurica complex. First, individuals (N = 7) ofthe newly described E. caurica samoensis ap-pear as a distinct lineage sister to individuals(N = 15) from the remainder of the Pacific andWestern Australia (E. caurica caurica). Fourgeographically structured haplotype clades arefound exclusively in the Western Indian Ocean.Erronea caurica dracaena is currently restrictedto the Seychelles based on sampling. Newlyadded individuals from East Africa and Mada-gascar form a haplotype clade that I recognizeas Erronea caurica elongata. Individuals of E.caurica quinquefasciata from the Red Sea, EastAfrica and Oman form the third monophyleticgroup. Finally, newly sequenced individualsfrom Masirah (N = 7) form a private haplotypeclade (E. caurica ssp. #1) sister to E. cauricaquinquefasciata. The final, newly added taxon(E. caurica ssp. #2) within the E. caurica com-plex is a clade (N = 18) that includes individu-als primarily from India, but with a fewindividuals from Masirah, Oman. This haplo-type clade is sister to the clade recognized pre-viously as E. caurica cf. derosa from the

Andaman Sea (Meyer, 2003). The Erroneacaurica complex and the associated E.cylindrica, E. ovum and E. errones species willbe more thoroughly addressed in another pa-per (Meyer, in prep.) as the group exhibits re-markable geographic structuring, polyphyly ofrecognized species (E. ovum), and evidenceof introgression based on nuclear markers.

Purpuradusta is well supported and containsfour newly added taxa that fall in expected re-lationships. The southeastern Polynesian en-demic species Purpuradusta oryzaeformis isdistinct and sister to P. minoridens that rangesthroughout the remainder of the westernIndoPacific. A single specimen of P. microdonfrom East Africa falls outside the haplotypeclade of other sampled individuals from thePacific basin (N = 5). This East African popula-tion is recognized as Purpuradusta microdonchrysalis. Two peripheral populations ofPurpuradusta fimbriata in the Pacific Basin areintroduced. First, Hawaiian populations of P.fimbriata are distinct (N = 7) and were previ-ously recognized as P. fimbriata waikikiensis;thus this name is resurrected as a valid entity.Second, individuals from the Marquesas arealso distinct genetically, consistent with thesubspecies designation of Lorenz (2002), P.fimbriata marquesana (N = 14). Both of thesePacific P. fimbriata subclades share a morerecent history with the widespread Pacific sub-species P. fimbriata unifasciata, as expected.

Two newly added species, “Erronea” barclayiand “Erronea” pulchella, come out as sisterspecies in phylogenetic analyses. Moreover,these two taxa appear as sister to Contradustain the most likely topology. Because of theseresults, and the poorly supported nature of theirrelationships, I tentatively place the two taxa inthe genus Contradusta, with the caveat thatthey may be removed with future data. Theseresults are somewhat surprising, particularlybecause “Contradusta” pulchella is thought tobe closely related to Erronea pyriformis be-cause of the darkly stained columellar denti-tion and overall conchological similarities. Thesister relationship between Contradustapulchella and C. barclayi is more acceptableas their divergence is deep, and the phyloge-netic affiliations of C. barclayi were more diffi-cult to predict based on morphological criteria.Another surprising result is the sister relation-ship between Notadusta martini and “Erronea”hungerfordi. Given these phylogenetic results,I tentatively place “Erronea” hungerfordi withinNotadusta, but with little confidence, althoughit is reasonably supported (73/4), and suspectthat it may be removed with more samples and


sequence data. Within the remaining Notadustacomplex, individuals of N. punctata trizonata(N = 9) form a monophyletic group sister to thePacific N. punctata punctata clade. Finally, inregards to Notadusta, “Notadusta” rabaulensiswas mentioned previously as a member ofErronea and “Notadusta” musumea asPalmulacypraea, further reducing the member-ship of Notadusta (Meyer, 2003).

The final four additions to the dataset fall intoMelicerona and Blasicrura. First, two taxa areadded to Melicerona. Samples of Meliceronalisteri melvilli (N = 5) from Queensland, Austra-lia, form a monophyletic group sister to the re-maining Melicerona taxa. (Two rostrate andmelanistic individuals interfinger among theother three haplotypes indicating that the tera-tology is likely driven by phenotypic responsesto environmental conditions rather than havinga genetic basis.) Samples of Melicerona felinafrom both Oman and East Africa form a mono-phyletic group, and because the haplotypesfrom the two regions interfinger, there is noevidence for a distinction between the subspe-cies M. felina felina and M. felina fabula. WithinBlasicrura, two taxa are added, based on thesequencing results. First, samples of Blasicrurasummersi, a Fijian and Tonga endemic, appearas a recently divergent sister to the also newlyincluded B. pallidula cf. vivia from AmericanSamoa. This clade is sister to the Melanesiansubspecies Blasicrura pallidula rhinoceros, asexpected based on geography. This resultingtopology indicates that the Blasicrura pallidulacomplex is paraphyletic.

DISCUSSION

The ultimate goal of this project is to constructa comprehensive phylogeny of cypraeid gas-tropods at the appropriate level for diversifica-tion studies. From a molecular perspective, allESUs presented are effectively equal units ofdiversity, whether they are currently recognizedas species, subspecies or some other level.There are some noted exceptions as OTUswere used on occasion that represented un-sorted or clinal variation within an ESU (e.g.,Erosaria miliaris/eburnea). However, on a gen-eral scale, each taxon shown in the phylogenies(Figs. 3−6) represents an independent evolu-tionary trajectory.

Because so much taxonomic information isavailable for cowries, it is informative to seehow molecular criteria compare with recog-nized taxonomic entities. The most recent com-pilation of the cowries is that of Lorenz (2002),

and I will use his checklist (pp. 250−291) as abenchmark for comparisons. Lorenz recog-nizes 232 species, of which I have sequenced210 (> 90%), and they are presented herein.The missing species are as follows:Nesiocypraea aenigma, N. lisetae, N. midway-ensis, Austrasiatica alexhuberti, Erosariaostergaardi, Zoila perlae, Lyncina camelopar-dis, L. joycae, Pustularia chiapponii, Cypra-eovula colligata, C. cruickshanki, C. immelmani,Palmadusta androyensis, P. johnsonorum,Austrasiatica deforgesi, Palmulacypraeaboucheti, P. omii, Eclogavena luchuana,Erronea (?) angioyorum, and E. nymphae. Se-quences from samples of both Purpuradustabarbieri and “Talostolida” rashleighana havebeen obtained, but were too late for inclusion inthese analyses. All missing species are rare,with small ranges located generally at the pe-riphery of their putative sister species based onconchological and anatomical characters. Ofthe 210 sequenced species, phylogenetic com-parisons and molecular criteria support all but15 (93%) as ESUs. The 15 recognized speciesnot supported by my criteria are discussed be-low. For Nucleolaria granulata, Monetariaobvelata, Erosaria eburnea, Zoila orientalis, Z.thersites, Luria controversa, L. gilvella,Notocypraea occidentalis, and Palmadustahumphreysii, multiple individuals were se-quenced and the haplotypes interfingeredwithin their closest relative. For the next sixspecies that I do not support, only a single in-dividual was sequenced, thus they may indeedrepresent a very young independent trajectory.However, when compared to the genetic diver-sity within their closest relative, the genetic dif-ference is unremarkable, and in someinstances, only a single mutation different fromputative conspecifics: Zonaria angelicae, Z.petitiana, Cypraeovula mikeharti, Bistolidabrevidentata, Cribrarula garciai, and C. taitae.

While genetic data are overall broadly con-sistent with taxa recognized at the specific level,the results are even more remarkable whencompared among taxa recognized at subspe-cific levels. Lorenz recognizes 260 taxa at thesubspecific level. Of those 260 subspecies, Ihave sequenced at least two individuals from160 in order to assess their validity. Molecularcriteria support 113 (> 70%) of these taxa aslegitimate ESUs. Moreover, sequence resultsindicate an additional 20 distinct ESUs notrecognized as subspecies by Lorenz (butsometimes mentioned as important varietiesor forms). A full listing of sampled taxa andtheir current ESU status as indicated by theprior criteria can be found at the Cowrie Ge-

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netic Database Project Website (http://www.flmnh.ufl.edu/cowries). The website in-cludes other information, such as localitiessampled, numbers of individuals for each taxon,and photographs of the specimens sequenced.

Overwhelming molecular support for tradition-ally recognized taxa, both at specific and sub-specific levels, is extremely encouraging. First,from a taxonomic standpoint, these molecularresults corroborate the excellent work done bycenturies of malacological researchers, at bothprofessional and amateur levels. Similar mo-lecular surveys of other diverse groups will pro-vide valuable comparisons in order to assesstaxonomic congruence (e.g., Jackson &Cheetham, 1990) and address concordant di-versification patterns. Second, from a molecu-lar perspective, sequence data provide asuitable, objective, relative metric for circum-scribing appropriate evolutionary units. Assum-ing rate constancy in the molecules (COI only,in prep.), molecular divergences can constrainthe tempo of diversification and assess the dis-tinctiveness of purported taxa. A growing bodyof molecular data across the diversity of lifeundoubtedly will provide insight to some of ourmost fundamental evolutionary questions.

ACKNOWLEDGEMENTS

An ever-growing number of individual andinstitutions have contributed and supported thisongoing research. Without their assistance, thework would not be possible. The following per-sons are recognized: Nonoy Alonzo, VicenteAzurin, Paul Barber, Don Barclay, Marty Beals,Victor Bonito, Philippe Bouchet, Michel Boutet,Roy Caldwell, Carlos Carvalho, Hank Chaney,John Chester, Peter Clarkson, Lori Bell Colin,Pat Colin, Allen Collins, Harry Conley, VinceCrayssac, Carolyn Cruz, Donald Dan, MartynDay, Bruno de Bruin, Helen deJode, John Earle,Andrew Edinger, Mark Erdmann, Melissa Frey,Michel Garcia, Bill Gibbs, Serge Gofas, TerryGosliner, Jeroen Goud, Robert Gourguet,Fabien Goutal, Paulo Granja, Kibata Mussa Haji,Jerry Harasewych, Itaru Hayami, Brian Hayes,Claus Hedegaard, Ed Heiman, Bert Hoeksema,John Hoover, John Jackson, Maurice Jay, ScottJohnson, Paul Kanner, Yasunori Kano, TomokiKase, Norbert Kayombo, Shigemitsu Kinjo, LisaKirkendale, Kitona Kombo Kitona, Utih Kukun,Senthil Kumar, Jean Paul Lefort, Bill Liltved,Hung-Chang Liu, Charlotte Lloyd, Felix Lorenz,Jr., Felix Lorenz, Sr., Larry Madrigal, MarleneMartinez, Gerald McCormack, MohammedMohammed, Hugh Morrison, Gowele Mtoka,

Mtumwa Mwadini, Peter Ng, Steve Norby,Shuichi Ohashi, Yoshihiro Omi, Ina Park, MarcelPin, Cory Pittman, Xavier Pochon, Matt Rich-mond, Raphael Ritson-Williams, Gonçalo Rosa,Gary Rosenberg, Teina Rongo, Fred Schroeder,Mike Severns, Pauline Severns, Hung-Long Shi,Brian Simison, Michael Small, John Starmer,Steve Tettlebach, David Touitou, Martin Wallace,Chia-Hsiang Wang, Dave Watts, Barry Wilson,Woody Woodman, Shu-Ho Wu. The followinginstitutions are acknowledged: Florida Museumof Natural History; University of California Mu-seum of Paleontology; Academy of Natural Sci-ences of Philadelphia; Bernice P. BishopMuseum, Honolulu, Hawaii; California Academyof Sciences; Institute of Marine Sciences, Zan-zibar; University of Dar es Salaam; Jackson-ville Shell Club; Musée National d’HistoireNaturelle, Paris, France; National Museum ofNatural History Naturalis, Leiden, The Nether-lands; Santa Barbara Museum of Natural His-tory; National Museum of Natural History; andSuganthi Devadason Marine Research Institute.I also would like to thank Felix Lorenz, Jr., forhis thoughtful comments, as well as the reviewsof four anonymous reviewers. Final decisionsand opinions are wholly mine.

This research has been financially supportedby the following NSF grants: DEB-9807316,DEB 0196049, and OCE-0221382.

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MEYER146

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161407

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(continues)

(continued)

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d localit

ies*

16S

C

OI

Mauritia m

auritiana

Lin

naeus, 1758

5, 19, 52, 70

AY

161474

AY

161707

Mauritia d

epre

ssa d

ispers

a

Schild

er

& S

child

er,

1939

10, 19

AY

161475

AY

161708

Mauritia d

epre

ssa d

epre

ssa

Gra

y, 1824

59, 64, 69

AY

161476

AY

161709

Mauritia g

rayana

Schild

er,

1930

2, 3

AY

161467

AY

161700

Mauritia e

gla

ntina

Dulc

os, 1833

32, 52, 59, 64

AY

161469

AY

161702

Mauritia h

istr

io

Gm

elin

, 1791

5, 9, 19

AY

161470

AY

161703

Mauritia a

rabic

a a

rabic

a

Lin

naeus, 1758

19, 21, 22, 23, 25, 29, 30, 32, 33, 52,

5

5, 56, 59, 61, 63, 64

AY

161465

AY

161698

17. M

au

riti

a a

rab

ica a

rab

ica (

Am

. S

am

oa)

67

AY

534367

AY

534444

Mauritia a

rabic

a a

sia

tica

Schild

er

& S

child

er,

1939

2, 11, 13, 15, 19, 23, 29, 30

AY

161466

AY

161699

Mauritia a

rabic

a im

manis

5, 7, 8

AY

161468

AY

161701

Mauritia m

aculif

era

(M

arq

uesas)

mart

yb

eals

i L

ore

nz, 2

00

2

71

AY

161472

AY

161705

Mauritia m

aculif

era

( N

. P

acific

) m

aculif

era

S

child

er,

1932

59, 66, 73

AY

161471

AY

161704

Mauritia m

aculif

era

(P

oly

nesia

) scin

data

L

ore

nz, 2

00

2

69, 70

AY

161473

AY

161706

18. B

ary

cyp

raea t

eu

lere

i C

azen

avett

e, 1846

3

AY

534368

AY

534445

19. B

ary

cyp

raea f

ult

on

i S

ow

erb

y III, 1903

106

AY

534369

AY

534446

Zoila

marg

inata

marg

inata

G

askoin

, 1849

43, 44, 45, 46

AY

161495

AY

161728

20. Z

oila m

arg

inata

kety

an

a

Rayb

au

di, 1

978

38, 39, 40

A

Y534370

AY

534447

Zoila

rosselli

C

otton, 1948

37, 38, 39, 40, 41, 42, 43

AY

161497

AY

161730

Zoila

venusta

S

ow

erb

y II, 1

846

39, 40, 41, 43

AY

161496

AY

161729

Zoila

elu

dens

Raybaudi, 1

991

35

AY

161499

AY

161732

Zoila

decip

iens

Sm

ith, 1880

33, 34

AY

161498

AY

161731

21. Z

oila m

ari

ellae

Rayb

au

di, 1

983

32 (

?)

AY

534371

AY

534448

Zoila

thers

ites

Gaskoin

, 1849

45, 46

AY

161503

AY

161736

Zoila

jeania

na s

herlyae

Raybaudi, 1

990

37

AY

161500

AY

161733

Zoila

jeania

na jeania

na

Cate

, 1968

36

AY

161501

AY

161734

Zoila

fri

en

dii

frie

ndii

Gra

y, 1831

39, 40, 41, 42, 43

AY

161502

AY

161735

Mura

cypra

ea m

us

Lin

naeus, 1758

81

AY

161491

AY

161724

Cypra

ea tig

ris (

India

n)

Lin

naeus, 1758

3, 5, 8, 9, 10, 14

AY

161489

AY

161722

Cypra

ea tig

ris (

Pacific

)

15, 18, 19, 23, 30, 32, 33, 52, 56, 57,

5

9, 60, 61, 62, 63, 64, 69, 70, 73

AY

161488

AY

161721

Cypra

ea p

anth

erina

Lig

htfoot, 1

786

1

AY

161490

AY

161723

(continues)

(continued)


A

ccessio

n #

OT

Us

Auth

ors

S

am

ple

d localit

ies*

16S

C

OI

Talp

aria talp

a

Lin

naeus, 1758

5, 30, 64

AY

161464

AY

161697

22. T

alp

ari

a e

xu

sta

S

ow

erb

y I, 1832

4

AY

534372

AY

534449

Luri

a lurid

a

Lin

naeus, 1758

84, 86, 87

AY

161462

AY

161695

Luria c

inere

a

Gm

elin

, 1791

83, 82, 105

AY

161461

AY

161694

Luria p

ulc

hra

G

ray, 1824

2

AY

161463

AY

161696

Luria tessella

ta

Sw

ain

son, 1822

73

AY

161460

AY

161693

Luri

a isab

ella

Lin

naeus, 1758

5, 8, 9, 20, 30, 65, 69, 73, 98

AY

161458

AY

161691

Luri

a isab

ella

mexic

an

a

Ste

arn

s, 1893

76, 79

AY

161459

AY

161692

Annepona m

ariae

Schild

er,

1927

59, 64, 70

AY

161443

AY

161676

Tro

na

ste

rcora

ria

Lin

naeus, 1758

87

AY

161441

AY

161674

Chely

cypra

ea testu

din

aria

Lin

naeus, 1758

5, 59, 64

AY

161442

AY

161675

Lyncin

a (

Austr

ocypra

ea)

reevei

Sow

erb

y I, 1832

41, 45

AY

161444

AY

161677

Lyncin

a (

cf. M

ioly

ncin

a)

port

eri

Cate

, 1966

97

AY

161448

AY

161681

Lyncin

a (

Are

sto

ides)

arg

us a

rgus

Lin

naeus, 1758

30, 59

AY

161445

AY

161678

23. L

yn

icn

a (

Are

sto

ide

s)

arg

us

c

on

tras

tria

ta

Perr

y, 1811

10

AY

534373

AY

534450

24. L

yn

cin

a (

Ca

llis

tocy

pra

ea)

bro

de

rip

ii

So

werb

y I, 1832

101

AY

534374

AY

534451

Lyncin

a (

Calli

sto

cypra

ea)

niv

osa

Bro

derip, 1827

13, 14

AY

161446

AY

161679

Lyncin

a (

Calli

sto

cypra

ea)

leucodon

Bro

derip, 1827

23, 57

AY

161447

AY

161680

Lyncin

a (

Calli

sto

cypra

ea)

aura

ntium

G

melin

, 1791

59, 54

AY

161449

AY

161682

Lyncin

a v

entr

iculu

s

Lam

arc

k, 1810

59, 69, 70

AY

161452

AY

161685

25. L

yn

cin

a v

en

tric

ulu

s (

Xm

as IO

)

19

AY

534375

AY

534452

Lyncin

a s

ulc

identa

ta

Gra

y, 1824

73

AY

161450

AY

161683

26. L

yn

cin

a k

uro

hara

i K

uro

da &

Hab

e, 1961

20

AY

534376

AY

534453

Lyncin

a s

child

ero

rum

Ir

edale

, 1939

69, 70, 73

AY

161451

AY

161684

Lyncin

a lynx

Lin

naeus, 1758

9, 30, 69, 5

A

Y161456

AY

161689

Lyncin

a v

itellu

s

Lin

naeus, 1758

5, 8, 30

AY

161457

AY

161690

Lyncin

a levia

than

Schild

er

& S

child

er,

1937

5, 20, 55, 59, 69, 73

AY

161453

AY

161686

Lyncin

a c

arn

eola

Lin

naeus, 1758

1, 9, 13, 20, 30, 52, 53, 55, 59, 69

AY

161454

AY

161687

Lyncin

a p

ropin

qua

Garr

ett, 1879

69, 71

AY

161455

AY

161688

Pustu

laria g

lobulu

s g

lobulu

s

Lin

naeus, 1758

59

AY

161507

AY

161740

Pustu

laria

glo

bulu

s b

revir

ostr

is

Schild

er

& S

child

er,

1938

5, 10

AY

161508

AY

161741

(continues)

(continued)

MEYER152

A

ccessio

n #

OT

Us

Auth

ors

S

am

ple

d localit

ies*

16S

C

OI

Pustu

laria b

istr

inota

ta b

istr

inota

ta

Schild

er

& S

child

er,

1937

27, 59, 64

AY

161504

AY

161737

27. P

ustu

lari

a b

istr

ino

tata

keelin

gen

sis

S

ch

ild

er

& S

ch

ild

er,

1940

98

AY

534377

AY

534454

28. P

ustu

lari

a b

istr

ino

tata

su

bla

evis

S

ch

ild

er

& S

ch

ild

er,

1938

69, 70

AY

534378

AY

534455

Pustu

laria

ma

uie

nsis

B

urg

ess, 1967

73

AY

161505

AY

161738

Pustu

laria

cic

erc

ula

Lin

naeus, 1758

5, 14, 59, 64, 69

AY

161506

AY

161739

Pustu

laria m

arg

arita

D

illw

yn, 1817

59

AY

161509

AY

161742

Neobern

aya s

padic

ea

Sw

ain

son, 1823

77

AY

161517

AY

161750

Pseudo

zonaria a

nnettae

Dall,

1909

78

AY

161518

AY

161751

Pseudo

zonaria a

rabic

ula

Lam

arc

k, 1810

79

AY

161519

AY

161752

29. P

se

ud

ozo

na

ria n

igro

pu

ncta

ta

Gra

y, 1828

90

AY

534379

AY

534456

Pse

udo

zona

ria

rob

ert

si

Hild

alg

o, 1906

80

AY

161520

AY

161753

30. S

ch

ild

eri

a a

ch

ati

dea

So

werb

y I, 1837

84

AY

534380

AY

534457

Zonaria s

anguin

ole

nta

G

melin

, 1791

87

AY

161512

AY

161745

31. Z

on

ari

a p

icta

G

ray, 1824

86

AY

534381

AY

534458

Zon

aria

zon

ari

a

Gm

elin

, 1791

87

AY

161513

AY

161746

Zon

aria

pyru

m s

en

eg

ale

nsis

S

child

er,

1928

87

AY

161510

AY

161743

Zonaria p

yru

m a

ngole

nsis

O

dhner,

1923

88

AY

161511

AY

161744

32. Z

on

ari

a p

yru

m a

ng

elicae

Clo

ver,

1974

87

AY

534382

AY

534459

33. N

oto

cyp

raea h

art

sm

ith

i S

ch

ild

er,

1967

96

AY

534383

AY

534460

Noto

cypra

ea p

iperita

G

ray, 1847

41, 43, 45, 48

AY

161524

AY

161757

Noto

cypra

ea p

ulic

aria

Reeve, 1846

41

AY

161525

AY

161758

Noto

cypra

ea c

om

pto

ni

Gra

y, 1847

43, 45, 47, 48, 49

AY

161521

AY

161754

Noto

cypra

ea a

ngusta

ta

Gm

elin

, 1791

47, 48

AY

161522

AY

161755

34. N

oto

cyp

raea a

ng

usta

ta (

Po

rt L

inco

ln)

45

AY

534384

AY

534461

Noto

cypra

ea d

ecliv

is

Sow

erb

y II, 1

870

47, 48

AY

161523

AY

161756

Cypra

eovula

connelli

Lilt

ved, 1983

102

AY

161527

AY

161760

Cypra

eovula

casta

nea

Hig

gin

s, 1868

6

AY

161526

AY

161759

Cypra

eovula

iuts

ui

Shik

am

a, 1974

102

AY

161528

AY

161761

Cypra

eovula

capensis

G

ray, 1828

102

AY

161530

AY

161763

35. C

yp

raeo

vu

la f

us

co

rub

ra

Sh

aw

, 1909

6

AY

534385

AY

534462

36. C

yp

raeo

vu

la f

us

co

den

tata

G

ray, 1825

6

AY

534386

AY

534463

Cypra

eovula

coro

nata

S

child

er,

1930

102

AY

161529

AY

161762

(continues)

(continued)


A

ccessio

n #

OT

Us

Auth

ors

S

am

ple

d localit

ies*

16S

C

OI

37. C

yp

raeo

vu

la m

ikeh

art

i L

ore

nz, 1985

6

AY

534387

AY

534464

38. C

yp

raeo

vu

la a

lgo

en

sis

G

ray, 1825

6

AY

534388

AY

534465

Cypra

eovula

edentu

la

Gra

y, 1825

102

AY

161531

AY

161764

Cypra

eovula

alfre

densis

S

child

er

& S

child

er,

1929

102

AY

161532

AY

161765

Palm

adusta

conta

min

ata

dis

tans

Schild

er

& S

child

er,

1938

5

AY

161555

AY

161789

Palm

adusta

conta

min

ata

conta

min

ata

S

ow

erb

y I, 1832

25, 32, 71

AY

161556

AY

161790

Palm

ad

usta

asellu

s c

f. b

itaenia

ta

Gere

t, 1

903

30, 63

AY

161543

AY

161777

Palm

ad

usta

asellu

s c

f. a

sellu

s

Lin

naeus, 1758

5

AY

161545

AY

161779

Palm

ad

usta

asellu

s c

f. v

espacea

Melv

ill, 1905

10, 23, 25

AY

161544

AY

161778

Palm

adusta

saula

e

Gaskoin

, 1843

13, 20

AY

161554

AY

161788

Palm

adusta

lentigin

osa

Gra

y, 1825

3, 11

AY

161557

AY

161791

Palm

ad

usta

hum

ph

reysii

Gra

y, 1825

55, 51

AY

161550

AY

161784

Palm

adusta

lute

a

Gm

elin

, 1791

23, 64

AY

161549

AY

161783

Palm

adusta

zic

zac z

iczac

Lin

naeus, 1758

25, 54

AY

161552

AY

161786

Palm

ad

usta

zic

za

c m

isella

P

err

y, 1811

5

AY

161553

AY

161787

Palm

adusta

dilu

culu

m

Reeve, 1845

5, 10

AY

161558

AY

161792

Palm

adusta

cla

ndestina p

asserina

Melv

ill, 1888

3, 5, 10, 8

AY

161548

AY

161782

39. P

alm

ad

usta

cla

nd

esti

na c

f. c

lan

desti

na

(An

da

man

) Lin

naeus, 1758

14

AY

534389

AY

534466

Palm

adusta

art

uffeli

Jousseaum

e, 1876

20

AY

161546

AY

161780

Palm

adusta

cla

ndestina c

landestina c

f.

candid

a

Pease, 1865

23, 30, 33, 38, 53, 55, 59, 64,

28

AY

161547

AY

161781

Bis

tolid

a s

tolid

a s

tolid

a

Lin

naeus, 1758

24, 32, 55, 59, 23, 51, 64

AY

161536

AY

161770

Bis

tolid

a s

tolid

a c

lavic

ola

M

elv

ill, 1888

5

AY

161534

AY

161768

40. B

isto

lid

a s

tolid

a d

iag

ues

Melv

ill, 1

888

10

AY

534390

AY

534467

Bis

tolid

a s

tolid

a r

ubig

inosa

Gm

elin

, 1791

30

AY

161535

AY

161769

Bis

tolid

a e

ryth

raeensis

S

ow

erb

y I, 1837

4

AY

161533

AY

161767

41. B

isto

lid

a o

wen

ii

So

werb

y I, 1837

100

AY

534391

AY

534468

Bis

tolid

a g

oodalli

i S

ow

erb

y I, 1832

69, 72

AY

161537

AY

161771

Bis

tolid

a h

irundo

Lin

naeus, 1758

14, 15, 52, 53, 55, 59, 12

AY

161539

AY

161773

Bis

tolid

a u

rsellu

s

Gm

elin

, 1791

25, 59

AY

161541

AY

161775

42. B

isto

lid

a u

rsellu

s (

An

dam

an

)

14

AY

534392

AY

534469

(continues)

(continued)

MEYER154

A

ccessio

n #

OT

Us

Auth

ors

S

am

ple

d localit

ies*

16S

C

OI

Bis

tolid

a k

ieneri k

ieneri

Hid

alg

o, 1906

5, 8, 9

AY

161542

AY

161776

Bis

tolid

a k

ieneri d

eprieste

ri A

S

child

er,

1933

29, 52

AY

161540

AY

161774

Bis

tolid

a k

ieneri d

eprieste

ri B

19, 28, 29

AY

161538

AY

161772

Ovatipsa c

hin

ensis

ch

ine

ns

is

Gm

elin

, 1791

5, 25, 29

AY

161559

AY

161793

43. O

vati

psa c

hin

en

sis

am

iges

Melv

ill &

Sta

nd

en

, 1904

38, 53, 59, 73, 23

AY

534393

AY

534470

Ovatipsa c

olo

ba

Melv

ill, 1888

14, 3

AY

161560

AY

161794

Talo

sto

lida tere

s

Gm

elin

, 1791

53, 59, 69, 98

AY

161561

AY

161795

Talo

sto

lida p

ellu

cens

Melv

ill, 1888

5, 14, 19, 30, 38, 59, 64, 73, 76, 80, 7

1

AY

161562

AY

161796

44. T

alo

sto

lid

a s

ub

tere

s

Wein

kau

ff, 1881

70

AY

534394

AY

534471

45. T

alo

sto

lid

a lati

or

Melv

ill, 1

888

73

AY

534395

AY

534472

Cribra

rula

gaskoin

i R

eeve, 1846

73

AY

161572

AY

161806

Cribra

rula

cath

olic

oru

m

Schild

er

& S

child

er,

1938

54

AY

161575

AY

161809

46. C

rib

raru

la t

ait

ae

Bu

rgess, 1993

67

AY

534396

AY

534473

Crib

raru

la c

um

ingii

Sow

erb

y I, 1832

69, 70

AY

161573

AY

161807

47. C

rib

raru

la g

arc

iai

Lo

ren

z &

Rain

es, 2001

75

A

Y534474

Cribra

rula

asta

ryi

Schild

er,

1971

71

AY

161574

AY

161808

Crib

raru

la c

rib

raria

com

ma

P

err

y, 1811

5

AY

161565

AY

161799

48. C

rib

raru

la p

ellis

serp

en

tis

Lo

ren

z, 1999

100

AY

534397

AY

534475

49. C

rib

raru

la e

so

ntr

op

ia f

ran

cesco

i L

ore

nz, 2002

100

AY

534398

AY

534476

Crib

raru

la c

ribe

llum

G

askoin

, 1849

7

AY

161563

AY

161797

Cribra

rula

esontr

opia

D

uclo

s, 1833

7

AY

161564

AY

161798

Crib

raru

la falla

x

Sm

ith, 1881

41

AY

161569

AY

161803

Cribra

rula

cribra

ria (

Andam

an)

au

str

alien

sis

L

ore

nz, 2

00

2

14, 38

AY

161567

AY

161801

50. C

rib

raru

la c

rib

rari

a c

f. a

balien

a

Lo

ren

z, 1989

3

AY

534399

AY

534477

Cribra

rula

gaspard

i B

iraghi, &

, N

icola

y, 1993

64

AY

161570

AY

161804

Crib

raru

la c

rib

raria

exm

outh

ensis

M

elv

ill, 1888

35

AY

161571

AY

161805

51. C

rib

raru

la e

xm

ou

then

sis

mag

nif

ica

Lo

ren

z, 2002

33

AY

534400

AY

534478

Crib

raru

la c

rib

raria

crib

raria

Lin

naeus, 1758

15, 24, 38, 52, 53, 55, 59, 23, 50, 98

AY

161566

AY

161800

52. C

rib

raru

la c

rib

rari

a m

elw

ard

i Ir

ed

ale

, 1930

50

AY

534401

AY

534479

Crib

raru

la c

rib

raria

rottn

este

nsis

R

aybaudi, 1

987

40

AY

161568

AY

161802

53. C

rib

raru

la c

rib

rari

a a

bro

lhen

sis

L

ore

nz, 2002

38

AY

534402

AY

534480

Nesio

cypra

ea

Au

str

asia

tica langfo

rdi

Kuro

da, 1938

93

AY

161516

AY

161749

(continues)

(continued)


A

ccessio

n #

OT

Us

Auth

ors

S

am

ple

d localit

ies*

16S

C

OI

Nesio

cypra

ea

Au

str

asia

tica h

irasei

Robert

s, 1913

25

AY

161514

AY

161747

Nesio

cypra

ea

Au

str

asia

tica s

akura

i H

abe, 1970

25

AY

161515

AY

161748

Palm

ula

cypra

ea k

ats

uae

Kuro

da, 1960

25

AY

161584

AY

161818

54. P

alm

ula

cyp

raea m

usu

mea

Ku

rod

a &

Hab

e, 1961

55

AY

534403

AY

534481

55. E

rro

nea

xan

tho

do

n

So

werb

y I, 1832

51

AY

534404

AY

534482

56. E

rro

nea p

allid

a

Gra

y, 1824

16

AY

534405

AY

534483

57. E

rro

nea v

red

en

bu

rgi

Sch

ild

er,

1927

104

AY

534406

AY

534484

58. E

rro

nea

rab

au

len

sis

S

ch

ild

er,

1964

95

AY

534407

AY

534485

59. E

rro

nea f

ern

an

do

i C

ate

, 1969

109

AY

534408

E

rronea (

Adusta

) onyx

Lin

naeus, 1758

3, 14, 20, 57, 11

AY

161608

AY

161842

Err

onea (

Adusta

) adusta

Lam

arc

k, 1810

5, 9

AY

161610

AY

161844

Err

on

ea

(A

dusta

) sub

vir

idis

sub

vir

idis

R

eeve, 1835

55, 51

AY

161612

AY

161846

Err

onea (

Adusta

) onyx m

ela

nesia

e

Schild

er,

1937

54

AY

161609

AY

161843

Err

onea (

Adusta

) subvirid

is d

ors

alis

S

child

er

& S

child

er,

1938

33, 38

AY

161611

AY

161845

60. E

rro

nea

pyri

form

is

Gra

y, 1824

107

AY

534409

AY

534486

Err

onea c

ylin

drica c

ylin

dri

ca

Born

, 1778

16, 24, 30, 33, 59

AY

161607

AY

161841

61. E

rro

nea c

ylin

dri

ca len

ella

Ired

ale

, 1939

55

AY

534410

AY

534487

62. E

rro

nea

ovu

m o

vu

m

Gm

elin

, 1791

16, 23

AY

534411

AY

534488

Err

on

ea

ovu

m p

ala

uensis

S

child

er

& S

child

er,

1938

57

AY

161601

AY

161835

Err

onea e

rrones

Lin

naeus, 1758

11, 14, 16, 30, 52, 55, 57, 58, 33, 53, 56

AY

161606

AY

161840

Err

on

ea

ovu

m c

hry

so

sto

ma

S

child

er,

1927

53, 54

AY

161600

AY

161834

Err

on

ea

cau

rica c

auri

ca

Lin

naeus, 1758

23, 30, 33, 38, 52, 53, 55, 64

AY

161602

AY

161836

Err

on

ea

cau

rica c

f. d

ero

sa

G

melin

, 1791

14, 11

AY

161603

AY

161837

Err

onea c

aurica d

racaena

Born

, 1778

9, 10

AY

161604

AY

161838

Err

on

ea

cau

rica q

uin

que

fascia

ta

Rodin

g, 1798

1, 3, 5

AY

161605

AY

161839

63. E

rro

nea c

au

rica e

lon

gata

P

err

y, 1811

5

AY

534412

AY

534489

64. E

rro

nea c

au

rica s

pp

. 1

3

AY

534413

AY

534490

65. E

rro

nea c

au

rica s

pp

. 2

3, 11

AY

534414

AY

534491

66. E

rro

nea c

au

rica s

am

oen

sis

L

ore

nz, 2002

55, 67

AY

534415

AY

534492

Purp

ura

dusta

serr

ulif

era

S

child

er

& S

child

er,

1938

69, 71

AY

161578

AY

161812

Purp

ura

dusta

min

oridens

Melv

ill, 1901

14, 32, 31, 98

AY

161577

AY

161811

67. P

urp

ura

du

sta

ory

zaefo

rmis

L

ore

nz &

Ste

rba, 1999

69, 70

AY

534416

AY

534493

(continues)

(continued)

MEYER156

A

ccessio

n #

OT

Us

Auth

ors

S

am

ple

d localit

ies*

16S

C

OI

Purp

ura

dusta

mic

rodon m

icro

do

n

Gra

y, 1828

52, 59, 56

AY

161576

AY

161810

68. P

urp

ura

du

sta

mic

rod

on

ch

rysalis

Kie

ner,

1843

5

AY

534417

AY

534494

Purp

ura

dusta

ham

mondae

Iredale

, 1939

23, 38, 98

AY

161579

AY

161813

Purp

ura

dusta

gra

cili

s n

ota

ta

Gill

, 1858

3, 5

AY

161581

AY

161815

Purp

ura

dusta

gra

cili

s g

racili

s

Gaskoin

, 1849

14, 15, 20, 28, 33, 59, 11, 51

AY

161580

AY

161814

Purp

ura

dusta

fim

briata

fim

briata

G

melin

, 1791

5, 9, 14

AY

161582

AY

161816

Purp

ura

dusta

fim

briata

unifascia

ta

Mig

hels

, 1845

30, 31, 59, 64, 69, 18, 19, 23, 61, 70

AY

161583

AY

161817

69. P

urp

ura

du

sta

fim

bri

ata

ma

rqu

esa

na

L

ore

nz, 2002

71

AY

534418

AY

534495

70. P

urp

ura

du

sta

fim

bri

ata

waik

ikie

nsis

S

ch

ild

er,

1933

73

AY

534419

AY

534496

Con

trad

usta

wa

lkeri

S

ow

erb

y I, 1832

14, 23, 38, 98

AY

161598

AY

161832

Contr

adusta

bre

geriana

Cro

sse, 1868

54, 92

AY

161599

AY

161833

71. C

on

trad

usta

? b

arc

layi

Reeve, 1857

106

AY

534420

AY

534497

72. C

on

trad

usta

? p

ulc

hella

Sw

ain

so

n, 1829

22

AY

534421

AY

534498

73. N

ota

du

sta

? h

un

gerf

ord

i S

ow

erb

y III, 1888

22

AY

534422

N

ota

dusta

mart

ini

Schepm

an, 1907

64

AY

161590

AY

161824

Nota

dusta

puncta

ta p

uncta

ta

Lin

naeus, 1771

23, 52, 53, 55, 59, 64

AY

161589

AY

161823

74. N

ota

du

sta

pu

ncta

ta t

rizo

nata

S

ow

erb

y II, 1

870

69, 70

AY

534423

AY

534499

Nota

dusta

puncta

ta (

Andam

an)

14

AY

161588

AY

161822

Nota

dusta

puncta

ta b

erinii

A

Dautz

enberg

, 1906

5

AY

161587

AY

161821

Nota

dusta

puncta

ta b

erinii

B

10

AY

161586

AY

161820

Melic

ero

na lis

teri

Gra

y, 1824

19, 30, 12

AY

161585

AY

161819

75. M

elicero

na lis

teri

melv

illi

Hid

alg

o, 1906

50

AY

534424

AY

534500

76. M

elicero

na f

elin

a

Gm

elin

, 1791

3, 5

AY

534425

AY

534501

Eclo

gavena d

ayritiana

Cate

, 1963

24

AY

161596

AY

161830

Eclo

gavena q

uadrim

acula

ta thie

lei

Schild

er

& S

child

er,

1938

33

AY

161591

AY

161825

Eclo

gavena q

uadrim

acula

ta q

uadrim

acula

ta

Gra

y, 1824

16, 24, 27, 32, 52, 57

AY

161592

AY

161826

Eclo

gavena c

oxeni

Cox, 1873

54

AY

161597

AY

161831

Bla

sic

rura

palli

dula

palli

dula

G

askoin

, 1849

15, 17

AY

161594

AY

161828

Bla

sic

rura

palli

dula

rhin

ocero

s

Souverb

ie, 1865

32, 52, 54

AY

161593

AY

161827

77. B

lasic

rura

pallid

ula

cf.

viv

ia

Ste

ad

man

& C

ott

on

, 1943

67

AY

534426

AY

534502

Bla

sic

rura

inte

rrupta

G

ray, 1824

14, 30, 12

AY

161595

AY

161829

78. B

lasic

rura

su

mm

ers

i S

ch

ild

er,

1958

92

AY

534427

AY

534503

(continued)

TOWARD COMPREHENSIVENESS: INCREASED MOLECULAR SAMPLING ... · TOWARD COMPREHENSIVENESS: INCREASED MOLECULAR SAMPLING ... Cowries (Gastropoda: Cypraeidae) ... and diversification patterns

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