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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
INCREASED MOLECULAR SAMPLING IN CYPRAEIDAE 129
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
INCREASED MOLECULAR SAMPLING IN CYPRAEIDAE 131
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
INCREASED MOLECULAR SAMPLING IN CYPRAEIDAE 133
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
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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
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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.
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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.
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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
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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-
MEYER144
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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Revised ms. accepted 19 January 2004
MEYER146
FIG
. 7
. S
am
ple
d lo
calit
ies
ind
ica
ted
fo
r lis
ted
ta
xa in
Ap
pe
nd
ix (
colu
mn
3).
INCREASED MOLECULAR SAMPLING IN CYPRAEIDAE 147
A
ccessio
n #
OT
Us
Auth
ors
S
am
ple
d localit
ies*
16S
C
OI
OU
TG
RO
UP
S
Jenneria p
ustu
lata
Lig
htfoot, 1
786
79
AY
161402
AY
161635
Pseudocypra
ea a
dam
sonii
Sow
erb
y I, 1832
5
AY
161403
AY
161636
1. P
seu
do
cyp
raea e
xq
uis
ita
Petu
ch
, 1979
23
AY
534351
AY
534428
Pedic
ula
ria p
acific
a
Pease, 1865
57, 59
A
Y161384
AY
161617
Cre
navolv
a tokuoi
Azum
a, 1989
25
AY
161390
AY
161623
Prio
novolv
a b
revis
S
ow
erb
y I, 1828
23
AY
161391
AY
161624
Serr
ato
volv
a d
ondani
Cate
, 1964
23
AY
161392
AY
161625
Phenacovolv
a tokio
i C
ate
, 1973
26
AY
161393
AY
161626
Cre
navolv
a c
f ro
sew
ate
ri
Cate
, 1973
25
AY
161394
AY
161627
2. V
olv
a v
olv
a
Lin
naeu
s, 1758
22
AY
534352
AY
534429
Phenacovolv
a w
eaveri
Cate
, 1973
25
AY
161395
AY
161628
3. P
rim
ovu
la c
on
cin
na
Ad
am
s &
Reeve, 1848
57
AY
534353
AY
534430
4. D
en
tio
vu
la t
akeo
i C
ate
& A
zu
ma, 1973
3
AY
534354
AY
534431
5. P
rosim
nia
sem
peri
W
ein
kau
ff, 1881
57
AY
534355
AY
534432
Adam
antia flo
rida
Kuro
da, 1958
25
AY
161396
AY
161629
Calp
urn
us v
err
ucosus
Lin
naeus, 1758
30
AY
161397
AY
161630
Calp
urn
us lacte
us
Lam
arc
k, 1810
59
AY
161398
AY
161631
Ovula
ovu
m
Lin
naeus, 1758
28
AY
161399
AY
161632
Cyphom
a g
ibbosum
Lin
naeus, 1758
83
AY
161400
AY
161633
Sim
nia
aequalis
S
ow
erb
y I, 1832
78
AY
161401
AY
161634
AP
PE
ND
IX 1
: S
am
ple
d T
axa
an
d S
up
ple
me
nta
ry D
ata
Ne
w t
axa
an
d s
eq
ue
nce
da
ta a
re n
um
be
red
an
d b
old
ed
. Ad
diti
on
al,
ne
wly
sa
mp
led
loca
litie
s fo
r ta
xa p
revi
ou
sly
rep
ort
ed
are
als
o b
old
ed
. C
ross
ou
tsa
re n
am
es
pre
vio
usl
y u
sed
in M
eye
r (2
00
3)
tha
t a
re c
orr
ect
ed
he
rein
be
cau
se o
f ta
xon
om
ic r
evi
sio
ns.
*fo
r th
e le
ge
nd
se
e F
ig.
7.
(continues)
MEYER148
A
ccessio
n #
OT
Us
Auth
ors
S
am
ple
d localit
ies*
16S
C
OI
ING
RO
UP
S
6. P
rop
ustu
lari
a s
uri
nam
en
sis
P
err
y, 1811
89
AY
534356
AY
534433
7. N
esio
cyp
raea t
era
mach
ii n
eo
cale
do
nic
a
Lo
ren
z, 2002
93
AY
534357
AY
534434
Ipsa c
hild
reni
Gra
y, 1825
59, 64
AY
161404
AY
161637
Cry
pto
cypra
ea d
illw
yni
Schild
er,
1922
59
AY
161411
AY
161644
Nucle
ola
ria n
ucle
us
Lin
naeus, 1758
5, 53, 64, 69
AY
161417
AY
161650
Nucle
ola
ria g
ranula
ta
Pease, 1862
73
AY
161418
AY
161651
Sta
phyla
ea lim
acin
a inte
rstincta
W
ood, 1828
5, 9
AY
161412
AY
161645
Sta
phyla
ea lim
acin
a lim
acin
a
Lam
arc
k, 1810
23, 30, 51
AY
161413
AY
161646
Sta
phyla
ea s
taphyla
ea laevig
ata
D
autz
enberg
, 1932
5, 8
AY
161414
AY
161647
Sta
phyla
ea s
taphyla
ea s
taphyla
ea
Lin
naeus, 1758
20, 51, 52, 53, 59
AY
161415
AY
161648
Sta
phyla
ea s
em
iplo
ta
Mig
hels
, 1845
73
AY
161416
AY
161649
Moneta
ria a
nnulu
s
Lin
naeus, 1758
3, 5, 8, 10, 30, 52, 68
AY
161405
AY
161638
Moneta
ria o
bvela
ta
Lam
arc
k, 1810
68, 69, 70
AY
161406
AY
161639
Moneta
ria m
oneta
Lin
naeus, 1758
5, 8, 30, 71, 59
AY
161407
AY
161640
Moneta
ria c
aputs
erp
entis c
aputo
phid
ii S
child
er,
1927
73
AY
161409
AY
161642
Moneta
ria c
aputs
erp
entis c
aputs
erp
entis
Lin
naeus, 1758
5, 19, 59, 66, 71, 91
A
Y161408
AY
161641
Moneta
ria c
aputd
raconis
M
elv
ill, 1888
75
AY
161410
AY
161643
Perissero
sa g
uttata
G
melin
, 1791
25
AY
161419
AY
161652
8. E
rosari
a m
arg
inalis
Dillw
yn
, 1827
5, 108
AY
534358
AY
534435
9. E
rosari
a c
itri
na
Gra
y, 1825
100, 101
AY
534359
AY
534436
Ero
saria h
elv
ola
helv
ola
Lin
naeus, 1758
5, 10, 30, 53, 59, 69, 70
AY
161429
AY
161662
Ero
saria h
elv
ola
haw
aiie
nsis
M
elv
ill, 1888
73
AY
161430
AY
161663
10. E
rosari
a h
elv
ola
cf.
callis
ta
Sh
aw
, 1909
71
AY
534360
AY
534437
Ero
saria turd
us
Lam
arc
k, 1810
1, 2
AY
161420
AY
161653
11. E
rosari
a irr
ora
ta
Gra
y, 1828
69, 70
AY
534361
AY
534438
Ero
saria a
lbugin
osa
Gra
y, 1825
79
AY
161427
AY
161660
Ero
saria p
ora
ria
Lin
naeus, 1758
19, 59, 69
AY
161428
AY
161661
Ero
saria b
eckii
Gaskoin
, 1836
59, 23, 31
AY
161425
AY
161658
12. E
rosari
a m
acan
dre
wi
So
werb
y II, 1
870
1
AY
534362
AY
534439
13. E
rosari
a e
ng
lert
i S
um
mers
& B
urg
ess, 1965
75
AY
534363
AY
534440
(continues)
(continued)
INCREASED MOLECULAR SAMPLING IN CYPRAEIDAE 149
A
ccessio
n #
OT
Us
Auth
ors
S
am
ple
d localit
ies*
16S
C
OI
Ero
saria k
ingae
Rehder
& W
ilson, 1975
74
AY
161424
AY
161657
Ero
saria thom
asi
Cro
sse, 1865
71
AY
161426
AY
161659
Ero
saria c
ern
ica
Sow
erb
y II, 1
870
38, 73
AY
161423
AY
161656
Ero
saria s
purc
a
Lin
naeus, 1758
84, 85, 86
AY
161421
AY
161654
Ero
sa
ria a
cic
ula
ris
Gm
elin
, 1791
83
AY
161422
AY
161655
Ero
saria labro
lineata
G
askoin
, 1849
52, 25
AY
161433
AY
161666
Ero
sa
ria b
oiv
inii
Kie
ner,
1843
20, 23
AY
161434
AY
161667
Ero
saria o
cella
ta
Lin
naeus, 1758
2, 12
AY
161431
AY
161664
Ero
saria g
angra
nosa
Dill
wyn, 1817
14
AY
161432
AY
161665
Ero
sa
ria lam
arc
kii
cf re
dim
ita
M
elv
ill, 1888
14
AY
161437
AY
161670
14. E
rosari
a lam
arc
kii lam
arc
kii
Gra
y, 1825
5
AY
534364
AY
534441
Ero
saria m
iliaris
Gm
elin
, 1791
15, 33
AY
161435
AY
161668
Ero
saria e
burn
ea
Barn
es, 1824
54, 55
AY
161436
AY
161669
Err
osaria n
ebrite
s
Melv
ill, 1888
1, 2
AY
161438
AY
161671
Ero
saria e
rosa (
India
n)
Lin
naeus, 1758
5, 8, 10, 14
AY
161440
AY
161673
Ero
saria e
rosa (
Pacific
) cf. c
hlo
rizan
s
Melv
ill, 1888
19, 30, 59, 69
AY
161439
AY
161672
Um
bili
a a
rmenia
ca
Verc
o, 1912
43, 44, 45
AY
161493
AY
161726
Um
bili
a h
esitata
Ir
edale
, 1916
48, 49
AY
161492
AY
161725
Um
bili
a c
ap
rico
rnic
a
Lore
nz, 1989
50
AY
161494
AY
161727
15. U
mb
ilia
cf.
peti
liro
str
is
Darr
ag
h, 2002
50
AY
534365
AY
534442
Macro
cypra
ea c
erv
inetta
Kie
ner,
1843
79
AY
161485
AY
161718
Macro
cypra
ea c
erv
us
Lin
naeus, 1771
83, 82
AY
161486
AY
161719
Macro
cypra
ea z
ebra
Lin
naeus, 1758
82, 83
AY
161487
AY
161720
Lep
ori
cyp
raea
va
len
tia
Perr
y, 1811
25
AY
161484
AY
161717
Leporicypra
ea m
appa r
osea
Gra
y, 1824
8
AY
161480
AY
161713
16. L
ep
ori
cyp
raea
ma
pp
a a
liw
ale
ns
is
Lo
ren
z, 2002
101
AY
534366
AY
534443
Leporicypra
ea m
appa m
appa
Lin
naeus, 1758
22, 23, 98
AY
161481
AY
161714
Leporicypra
ea g
eogra
phic
a
Schild
er
& S
child
er,
1933
5, 10, 14, 32, 54, 56, 57, 98
AY
161482
AY
161715
Leporicypra
ea m
appa p
anery
thra
vir
idis
M
elv
ill, 1888
; K
enyon, 1902
55, 57, 59, 64
AY
161479
AY
161712
Leporicypra
ea m
appa v
irid
is a
dm
irab
ilis
K
enyon, 1902
; Lore
nz, 2002
71
AY
161483
AY
161716
Mauritia s
curr
a indic
a
Gm
elin
, 1791
30, 59, 71
AY
161477
AY
161710
Mauritia s
curr
a s
curr
a
Gm
elin
, 1791
5, 10
AY
161478
AY
161711
(continues)
(continued)
MEYER150
A
ccessio
n #
OT
Us
Auth
ors
S
am
ple
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)
INCREASED MOLECULAR SAMPLING IN CYPRAEIDAE 151
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)
INCREASED MOLECULAR SAMPLING IN CYPRAEIDAE 153
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)
INCREASED MOLECULAR SAMPLING IN CYPRAEIDAE 155
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)