Puillandre, N., Duda, T.F., Jr., Meyer, C

Journal of Molluscan Studies Advance Access published 5 September 2014

Journal of The Malacological Society of London Molluscan Studies

Journal of Molluscan Studies (2014) 1–23. doi:10.1093/mollus/eyu055

One, four or 100 genera? A new classiﬁcation of the cone snails

N. Puillandre1,T.F.Duda2,C.Meyer3, B. M. Olivera4 and P. Bouchet5 1Museum National d’Histoire Naturelle, Departement Systematique et Evolution, ISyEB Institut (UMR 7205 CNRS/UPMC/MNHN/EPHE), 43, Rue Cuvier, Paris 75231, France; 2Department of Ecology and Evolutionary Biology and Museum of Zoology, University of Michigan, 1109 Geddes Avenue, Ann Arbor, MI 48109, USA; 3Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA; 4Department of Biology, University of Utah, 257 South 1400 East, Salt Lake City, UT 84112, USA; and 5Museum National d’Histoire Naturelle, Departement Systematique et Evolution, ISyEB Institut (UMR 7205 CNRS/UPMC/MNHN/EPHE), 55, Rue Buffon, Paris 75231, France Downloaded from

Correspondence: N. Puillandre; e-mail: [email protected] (Received 7 August 2013; accepted 6 June 2014)

ABSTRACT http://mollus.oxfordjournals.org/ We present a new classification for the genus Conus sensu lato (family Conidae), based on molecular phylogenetic analyses of 329 species. This classification departs from both the traditional classification in only one genus and from a recently proposed shell- and radula-based classification scheme that separates members of this group into five families and 115 genera. Roughly 140 genus-group names are available for Recent cone snails. We propose to place all cone snails within a single family (Conidae) containing four genera—Conus, Conasprella, Profundiconus and Californiconus (with Conus alone encompassing about 85% of known species)—based on the clear separation of cone snails into four distinct and well-supported groups/ lineages in molecular phylogenetic analyses. Within Conus and Conasprella, we recognize 57 and 11 subgenera, respectively, that represent well-supported subgroupings within these genera, which we interpret as by guest on December 3, 2014 evidence of intrageneric distinctiveness. We allocate the 803 Recent species of Conidae listed as valid in the World Register of Marine Species into these four genera and 71 subgenera, with an estimate of the confi- dence for placement of species in these taxonomic categories based on whether molecular or radula and/or shell data were used in these determinations. Our proposed classification effectively departs from previous schemes by (1) limiting the number of accepted genera, (2) retaining the majority of species within the genus Conus and (3) assigning members of these genera to species groups/subgenera to enable the effective communication of these groups, all of which we hope will encourage acceptance of this scheme.

INTRODUCTION After more than two centuries of naming and overnaming, the species-level systematics of cones is undergoing a re- Cone snails are members of a hyperdiverse group of marine gas- appraisal based, among others, on molecular characters, and tropods that are well known and appreciated for their history of this is leading to a more stable taxonomy. Conversely, the rapid diversification, their tremendously diverse and complex supraspecific classification of the cone snails has in the last 20 venoms, and the elegance of their shells. They include such em- years become more unstable than ever. A plethora of nominal blematic species as the once mythical ‘Glory-of-the-Sea’ Cone (sub)genera reflect subtle differences in shell form and radular (Conus gloriamaris) and the deadly Geography Cone (Conus geo- morphology, but these are obscuring phylogenetic relation- graphus). From 30 valid species known to Linnaeus, the current ships between terminal taxa. While toxinologists are longing accepted number of species stands at 803 (WoRMS, 2014). for a predictive classification that can be used as a roadmap These live in the tropical and warm temperate seas of the world, for bioprospecting (Olivera, 2006; Puillandre & Holford, from the intertidal to depths of almost 1,000 m. Like most 2010), the two genus-level classifications of cones proposed in species in the superfamily Conoidea, cones are characterized by the last 25 years (da Motta, 1991; Tucker & Tenorio, 2009, the possession of a venom gland and a highly modified radular 2013) have remained ignored outside the world of cone-shell tooth that they use as a harpoon to inject venom into their prey. collectors. Toxins included in these venoms are being studied for the possi- Whereas the general and characteristic shape of the shell has bilities they offer to understand the mechanisms underlying the empirically brought together all cone snails in what for two cen- evolution of species interactions, ecological diversification and turies has been called the family Conidae, the relationships the functioning of the nervous system, and also for potential between the species within Conidae, and in particular their therapeutic applications. eventual classification in genera and subgenera, is still debated.

The first modern divisions of the family Conidae recognized one Identifying groups to be named family, 14 groups “which may even represent subfamilies”, and 29 genera (Cotton, 1945: 231), two subfamilies and eight genera As far as possible, only well supported (bootstrap probability . . (Walls, 1978) and eight genera and 60 subgenera (da Motta, 90%; Bayesian posterior probability 0.95) clades (and 1991). These classifications were based solely on shell characters, single-species lineages) were linked to names. When several whereas Tucker & Tenorio’s (2009, 2013) completely novel clas- alternatives were possible (e.g. one supported clade that includes sification was based on shell as well as radular characters. These two supported clades, both with available names attributable to authors divided cone snails into five families and 89 genera, them), other characters and properties, such as shell morphology, while their restricted family Conidae was itself subdivided into type of prey, bathymetric and/or geographical distribution, were four subfamilies. The robustness of this classification is question- considered in order to identify the most appropriate grouping to able, when radular characters have been found to be a poor minimize within-group variability. Overall, a conservative ap- phylogenetic marker in other conoidean taxa (Castelin et al., proach was adopted, to minimize the number of supraspecific 2012; Kantor & Puillandre, 2012). In fact, cone snails, or Conus taxa, both named and unnamed. In two cases (Pyruconus and sensu lato, is in itself a concept with blurred limits. Depending on Cylinder) we attributed a name to a group we recognized to be the authors, some marginal species may or may not be included non-monophyletic (although this non-monophyly is not sup- in cone snails. This is the case of the Cryptoconidae sensu Tucker ported), in order to avoid having to establish new names for & Tenorio (2009), treated by some authors as cone snails (e.g. many small clades. Such polyphyletic, but morphologically con- Cossmann, 1896), while excluded by others (Tucker & Tenorio, sistent, genera may correspond to grades and in future each of 2009). DNA data indicate that Genota, a genus sometimes the included clades may be shown to deserve its own name. included in this family, may be more closely related to the family Borsoniidae (Puillandre et al., 2011). Similarly, Naming of groups Downloaded from Benthofascis was excluded from the Conidae by Puillandre et al. (2011). Consequently, the present study will not consider As far as possible, a genus-group name (i.e. a genus or subgenus) Cryptoconidae and Benthofascis as members of the ingroup, but was applied based on the position of its type species in the tree. will focus on the Conidae sensu Bouchet et al. (2011), i.e. If the type species of a nominal genus or subgenus had not been Conidae, Conilithidae and Taranteconidae sensu Tucker & sequenced, application of the name was determined by reference to the morphologically most similar species used in the molecu- Tenorio (2009). http://mollus.oxfordjournals.org/ In the last decade, published molecular phylogenies (listed by lar analysis. If more than one name was applicable for a clade, Puillandre et al., 2014) have suggested that cone snails actually the valid name was determined by the rule of priority. In the encompass very deep lineages that diverged at least 33 Ma classification below, OD refers to the fixation of type species by (Duda & Kohn, 2005), the three main ones being the single original designation, SD by subsequent designation and M by species Conus californicus, sister-group to all the other cones, the monotypy (as defined by ICZN, 1999: Art. 68, 69, respectively). ‘Small Major Clade’ of Duda & Kohn (2005) (corresponding to the family Conilithidae of Tucker & Tenorio, 2009) and the Ranking of groups ‘Large Major Clade’ of Duda & Kohn, 2005 (corresponding to the family Conidae of Tucker & Tenorio, 2009). Furthermore, Tucker & Tenorio (2009, 2013) attributed family-rank names to Espiritu et al. (2001) defined 21 molecular groups within the the main lineages identified by molecular phylogenies (i.e. the Large Major Clade. None of these molecular phylogenies was Small and Large Major Clades) and genus-rank names to by guest on December 3, 2014 turned into a formal classification of cone snails. However, groups of species that roughly correspond to the same level of Bandyopadhyay et al. (2008) and a few others after him (e.g. resolution as the 21 groups defined by Espiritu et al. (2001). Kraus et al., 2011, 2012) did apply the name Conasprella to the However, contrary to the first two steps of the process, which are Small Major Clade, thus leaving the name Conus for the Large based on objective criteria (e.g. monophyly of the taxa) and no- Major Clade. Several other generic or subgeneric names have menclatural rules (e.g. position of type species), the attribution also been used sporadically (e.g. Jimenez, Olivera & Teichert, of ranks to the named clades is subjective. Taxonomists generally 2007; Holford et al., 2009; Puillandre, Watkins & Olivera, 2010). place two species in two different genera or two different families Meanwhile, new names have continued to be established to this (or any other rank) based on a certain level of divergence, but day, with no less than 127 genus-group names available to clas- the threshold of divergence at which a given rank is recognized sify cones. varies greatly depending on the taxa, the characters involved Puillandre et al. (2014) recently published a phylogeny of the and who is making the classification (Avise & Liu, 2011). Even cone snails based on 330 species and sequences of three mito- for a given taxon and character (e.g. a nucleotide sequence), it is chondrial gene regions. In the present paper we utilized this mo- difficult to provide an objective criterion, since ranks above the lecular phylogeny as a foundation to establish a new genus- and species level are not defined by any biological attribute or prop- subgenus-level classification of the Conidae, with four genera erty that could define a threshold. (For the rank of species, on (Californiconus, Profundiconus, Conasprella and Conus) and 71 sub- the other hand, objective criteria for delimitation have been pro- genera (Fig. 1). We also tentatively allocate all cone snail species posed, as amply developed in the literature on integrative tax- currently considered as valid in WoRMS, but not represented in onomy, e.g. De Queiroz, 1998; Samadi & Barberousse, 2006.) the molecular phylogeny, to genera and subgenera based on Accepting that ranking is to a large extent a matter of opinion, their morphological characters. we also took into consideration prevailing usage. In the vast literature dealing with conotoxins, ranking has been very conservative, with Conidae being the only accepted family-level taxon. We have also been guided by dated molecular phylogenies of MATERIAL AND METHODS other marine gastropod clades. For example, the mean age of Turning the phylogeny into a classification the most recent common ancestors of each monophyletic genus of Muricidae (Neogastropoda) is about 20 Ma, with a minimum Proposing a classification based on a phylogeny mainly consists of 9.5 Ma and a maximum of 42.2 Ma, and the age of the of (1) identifying the groups that will be named; (2) attributing subfamily Rapaninae is estimated at about 61 Ma (Claremont available names and, if necessary, establishing new ones, for the et al., 2013: fig. 2). The most recent common ancestors of the groups identified in (1); and (3) ranking these names. bursids Tutufa and Bursa (Littorinimorpha: Tonnoidea) are

2 CLASSIFICATION OF CONE SNAILS Downloaded from http://mollus.oxfordjournals.org/ by guest on December 3, 2014

Figure 1. Simplified version of the Bayesian tree based on the concatenation of sequences of three mitochondrial gene regions (COI, 16S, 12S) and published by Puillandre et al. (2014: fig. 2) showing the proposed classification. Genera and subgenera with multiple species are reduced to triangles, whose lengths are proportional to the branch lenghts. Posterior probabilities (.0.95) are shown for each node. Only (sub)genera with at least one sequenced representative are figured.

3 N. PUILLANDRE ET AL. estimated to be about 26 and 35 Ma, respectively (Castelin et al., SYSTEMATICS 2012: fig. 7). The ages of the genera Littoraria and Echinolittorina (Littorinimorpha: Littorinidae) and of Turbo (Vetigastropoda: Superfamily CONOIDEA Fleming, 1822 Turbinidae) are estimated to be respectively 55 Ma (Reid, Dyal Family Conidae Fleming, 1822 & Williams, 2012), 50 Ma (Reid et al., 2012) and 60–117 Ma (Williams, 2007; Williams & Duda, 2008), respectively. To place [Synonyms: Conilithidae Tucker & Tenorio, 2009, n. syn. these numbers in perspective, the oldest fossils attributed with Taranteconidae Tucker & Tenorio, 2009, n. syn. Puncticuliinae certainty to cone snails are 55 Ma (Duda & Kohn, 2005), the Tucker & Tenorio, 2009, n. syn.] minimum age for the divergence between the two main clades of cones is estimated to be 33 Ma (Duda & Kohn, 2005) and the species diversification within each of these two main clades is Genus Californiconus Tucker & Tenorio, 2009 supposed to have started between 15 and 20 Ma (Duda & Type species: Conus californicus Reeve, 1844; OD Palumbi, 1999). The main lineages within cone snails are thus roughly of the same age, or even younger, than the genera as Included species: defined in other groups of caenogastropods. In an attempt to californicus (Reeve, 1844) assign ranks that are roughly comparable with those used for other gastropod taxa in terms of group age, we chose to rank the Remarks: Conus californicus has always been considered a unique main lineages of cone snails at the level of genus. With this ap- species within cone snails, because of its molecular (including proach, all cone snails are placed in a single family (Conidae) toxicological: Biggs et al., 2010; Elliger et al., 2011) and morpho- and most (Large Major Clade) remain in the genus Conus (con- logical singularities and also because of its diet, since it is able to prey indifferently on fish, molluscs and worms (Kohn, 1966). sistent with prevailing usage of this name). As stated above, Downloaded from based on the molecular clock calibrations of Duda & Kohn (2005), the lineages that ultimately gave rise to these genera pre- sumably separated from each other at least 33 Ma. We further Genus Profundiconus Kuroda, 1956 identified well-supported groupings within these genera and Type species: Chelyconus profundorum Kuroda, 1956; OD assigned the members of these groups to subgenera. Except for [Synonym: Lizaconus da Motta, 1991; type species: Asprella the difference in assigned rank, these subgenera are essentially http://mollus.oxfordjournals.org/ comparable with the generic-level groups proposed by Tucker & teramachii Kuroda, 1956; OD] Tenorio (2009, 2013). Included species: cakobaui (Moolenbeek, Ro¨ckel pacificus (Moolenbeek & & Bouchet, 2008) Ro¨ckel, 1996) n. comb. Allocation of species to genus-level groups dondani (Kosuge, 1981) n. comb. profundorum (Kuroda, emersoni (G. D. Hanna, 1963) 1956) All the 803 species of Conidae listed as valid in WoRMS (2014) frausseni (Tenorio & Poppe, scopulicola (Okutani, 1972) were allocated to genera and subgenera, with two levels of confi- 2004) smirna (Bartsch & Rehder, dence. Those printed in bold font were placed in the classifica- ikedai (Ninomiya, 1987) 1943) tion based on the DNA sequences of Puillandre et al. (2014).

jeanmartini (G. Raybaudi stahlschmidti (Tenorio & by guest on December 3, 2014 The others were classified based on their shell and/or radula Massilia, 1992) Tucker, 2014) characters, following Tucker & Tenorio (2013) and, for species kanakinus (Richard, 1983) n. teramachii (Kuroda, 1956) not included or considered as synonyms by Tucker & Tenorio comb. tuberculosus (Tomlin, 1937) n. (2013), following Petuch (2013) or the advice of M. Tenorio lani (Crandall, 1979) comb. (personal communication). loyaltiensis (Ro¨ckel & vaubani (Ro¨ckel, Richard & Species sometimes referred to as ‘cone snails’ but allocated to Moolenbeek, 1995) Moolenbeek, 1995) Artemidiconus (Conorbidae; Tucker & Tenorio, 2009; Bouchet et al., 2011), Benthofascis (Conorbidae; Tucker & Tenorio, 2009; Bouchet et al., 2011), Genota (Cryptoconidae in Tucker & Tenorio, Remarks: Puillandre et al. (2014) did not sequence the type 2009; or Borsoniidae in Bouchet et al., 2011)andGenotina species of Profundiconus; their sequenced material was identified (Cryptoconidae in Tucker & Tenorio, 2009; or Mangeliidae in as P. aff. profundorum. However, Tucker & Tenorio (2009, 2013) Bouchet et al., 2011) were excluded. Within Conorbidae, only found Profundiconus to be a morphologically well supported Benthofascis lozoueti has been sequenced and molecular analysis group and we are thus confident in applying this name to the indicates that the family is separate from Conidae (Puillandre clade containing P. aff. profundorum. et al., 2011). However, B. lozoueti is the only conorbid species that does not resorb the inner shell walls (Tucker, Tenorio & Stahlschmidt, 2011). It thus cannot be excluded that the other Genus Conasprella Thiele, 1929 Conorbidae species—which resorb them—may in fact not be con- familial. Likewise, as indicated above, molecular data place Genota Subgenus Conasprella Thiele, 1929 in the Borsoniidae (Puillandre et al., 2011) and this clade is not Type species: Conus pagoda Kiener, 1847; SD, Tucker & Tenorio further discussed here. Fossil taxa (Hemiconus Cossmann, 1889; (2009: 140) (under ICZN, 1999: Art. 70.3.2). Cryptoconus Koenen, 1867; Conorbis Swainson, 1840; Conilithes Swainson, 1840; Eoconus Tucker & Tenorio, 2009 and Plagioconus [Synonym: Duodenticonus Tucker & Tenorio 2013; type species: Tucker & Tenorio, 2009) are not discussed either. Consequently, Asprella memiae Habe & Kosuge, 1970; OD; n. syn.] only the Conidae, Conilithidae and Taranteconidae (sensu Tucker & Tenorio, 2009) are discussed below, i.e. the cone snails Included species: as defined by Bouchet et al. (2011). aphrodite (Petuch, 1979) boholensis (Petuch, 1979) The classification is summarized as a list of the valid Recent articulata (G.B. Sowerby II, n. comb. species with their genus-level assignment (Appendix Table 1) 1873) boucheti (Richard, 1983) and as a list of nominal genus-group names with their current baileyi (Ro¨ckel & da Motta, n. comb. status (Appendix Table 2). 1979) bozzettii (Lauer, 1991)

4 CLASSIFICATION OF CONE SNAILS eucoronata (G.B. Sowerby memiae (Habe & Kosuge, Included species: III, 1903) 1970) n. comb. aculeiformis (Reeve, 1844) joliveti (Moolenbeek, Ro¨ckel eugrammata (Bartsch & otohimeae (Kuroda & Itoˆ, n. comb. & Bouchet, 2008) n. comb. Rehder, 1943) 1961) n. comb. comatosa (Pilsbry, 1904) coriolisi (Ro¨ckel, Richard & gordyi (Ro¨ckel & Bondarev, pagoda (Kiener, 1847) n. comb. Moolenbeek, 1995) n. comb. 2000) wakayamaensis (Kuroda, dictator (Melvill, 1898) n. comb. lizarum (Raybaudi Massilia & lorenzi Monnier & Limpalae¨r, 1956) dieteri (Moolenbeek, da Motta, 1992) n. comb. 2012 Zandbergen & Bouchet, longurionis (Kiener, 1847) 2008) n. comb. n. comb. elegans (G. B. Sowerby III, orbignyi (Audouin, 1831) Subgenus Boucheticonus Tucker & Tenorio, 2013 1895) n. comb. n. comb. Type species: Conus alisi Moolenbeek, Ro¨ckel & Richard, elokismenos (Kilburn, 1975) pepeiu (Moolenbeek, 1995; OD n. comb. Zandbergen & Bouchet, ﬁjiensis (Moolenbeek, Ro¨ckel & 2008) n. comb. Included species: Bouchet, 2008) n. comb. pseudorbignyi (Ro¨ckel & alisi (Moolenbeek, Ro¨ckel & pseudokimioi da Motta & guidopoppei (G. Raybaudi Lan, 1981) n. comb. Richard, 1995) n. comb. Martin, 1982 n. comb. Massilia, 2005) n. comb. saecularis (Melvill, 1898) kimioi (Habe, 1965) n. comb. hypochlorus (Tomlin, 1937) n. comb. n. comb. stocki (Coomans & Remarks: The species C. kimioi was placed in the subgenus hopwoodi (Tomlin, 1937) Moolenbeek, 1990) n. comb. Boucheticonus (following Tucker & Tenorio 2013), even if the cor- n. comb. tiki (Moolenbeek, Zandbergen Downloaded from responding clade was not highly supported. ichinoseana (Kuroda, 1956) & Bouchet, 2008) n. comb. n. comb. traversiana (E. A. Smith, 1875) insculpta (Kiener, 1847) n. comb. n. comb. Subgenus Coltroconus Petuch, 2013 lentiginosa (Reeve, 1844) viminea (Reeve, 1849) Type species: Conus iansa Petuch, 1979; OD n. comb. n. comb.

Included species: http://mollus.oxfordjournals.org/ henriquei (Petuch & iansa (Petuch, 1979) n. comb. Subgenus Kohniconus Tucker & Tenorio, 2009 R. F. Myers, 2014) n. comb. Type species: Conus emarginatus Reeve, 1844; OD Included species: Subgenus Dalliconus Tucker & Tenorio, 2009 arcuata (Broderip & janowskyae (Tucker & Tenorio, G. B. Sowerby I, 1829) 2011) n. comb. Type species: Conus mcgintyi Pilsbry, 1955; OD n. comb. kohni (McLean & Nybakken, Included species: centurio (Born, 1778) n. comb. 1979) n. comb. armiger (Crosse, 1858) n. comb. mcgintyi (Pilsbry, 1955) emarginata (Reeve, 1844) rachelae (Petuch, 1988) bajanensis (Nowell-Usticke, n. comb. n. comb. n. comb. by guest on December 3, 2014 1968) n. comb. pacei (Petuch, 1987) n. comb. fenzani (Petuch & Sargent, scaripha (Dall, 1910) guyanensis (van Mol, 1973) rainesae (McGinty, 1953) 2011) n. comb. n. comb. n. comb. n. comb. Remarks: Tucker & Tenorio (2009) included both Conasprella lenhilli (Cargile, 1998) n. comb. roberti (Richard, 2009) delesserti and Conasprella arcuata in Kohniconus. In our analysis, the mazei (Deshayes, 1874) n. comb. two species do not cluster together. The name Kohniconus could n. comb. sauros (Garcia, 2006) n. comb. have been applied to the lineage that includes the species C. delessertii, but because of the closer morphological resem- Subgenus Endemoconus Iredale, 1931 blance of Conus emarginatus with C. arcuata, we have applied Type species: Conus howelli Iredale, 1929; OD Kohniconus to the lineage that includes C. arcuata. [Synonyms: Kermasprella Powell, 1958; type species: Conus raoulensis Powell, 1958; OD. Yeddoconus Tucker & Tenorio, 2009; type Subgenus Lilliconus G. Raybaudi Massilia, 1994 species: Conus sieboldii Reeve, 1848; OD] Type species: Conus biraghii G. Raybaudi Massilia, 1992; OD Included species: Included species: grohi (Tenorio & Poppe, 2004) sieboldii (Reeve, 1848) biraghii (G. Raybaudi Massilia, sagei (Korn & G. Raybaudi n. comb. n. comb. 1992) n. comb. Massilia, 1993) n. comb. howelli (Iredale, 1929) n. comb. somalica (Bozzetti, 2013) ione (Fulton, 1938) n. comb. n. comb. Subgenus Parviconus Cotton & Godfrey, 1932 raoulensis (Powell, 1958) spiroﬁlis (Habe & Kosuge, n. comb. 1970) n. comb. Type species: Conus rutilus Menke, 1843; OD Included species: rutila (Menke, 1843) n. comb. Subgenus Fusiconus da Motta, 1991 Type species: Conus longurionis Kiener, 18547; OD Subgenus Pseudoconorbis Tucker & Tenorio, 2009 [Synonyms: Bathyconus Tucker & Tenorio, 2009; type species: Type species: Conorbis coromandelicus E. A. Smith, 1894; OD Conus orbignyi Audouin, 1831; OD; n. syn. Fumiconus da Motta, 1991; type species: Conus traversianus E.A. Smith, 1875, Included species: OD; n. syn. Viminiconus Tucker & Tenorio, 2009; type species: coromandelica (E. A. Smith, traceyi (Tucker & Stahlschmidt, Conus vimineus Reeve, 1849; OD; n. syn.] 1894) n. comb. 2010) n. comb.

5 N. PUILLANDRE ET AL.

Subgenus Ximeniconus Emerson & Old, 1962 Subgenus Asprella Schaufuss, 1869 Type species: Conus ximenes Gray, 1839; OD Type species: Conus asper Lamarck, 1810, by typification of replaced name. Asprella was established as a substitute name for [Synonyms: Globiconus Tucker & Tenorio, 2009; type species: Cylindrella Swainson, 1840 (see below). Conus asper was not Conus tornatus G.B. Sowerby I, 1833; OD; n. syn. among the 11 species included in Asprella by Schaufuss (1869: Jaspidiconus Petuch, 2003; type species: Conus jaspideus 43–44), and Wenz (1940) cited Conus sulcatus Bruguie` re, 1792 (a Gmelin, 1791; OD; n. syn. Perplexiconus Tucker & Tenorio, species originally included by Schaufuss) as the type species; 2009; type species: Conus perplexus G. B. Sowerby II, 1857; Conus sulcatus and C. asper are subjective synonyms. OD; n. syn.] [Synonyms: Cylindrella Swainson, 1840; type species: Conus asper Included species: Lamarck, 1810; subjective synonym of Conus sulcatus Hwass, alexandremonteiroi (Cossignani, ogum (Petuch & R. F. Myers, 1792; M. Cylindrella Swainson, 1840 (Conidae), is a homonym of 2014) n. comb. 2014) n. comb. Cylindrella Swainson, 1840 (Urocoptidae) and has been placed allamandi (Petuch, 2013) perplexa (G. B. Sowerby II, on the Official Index of Rejected and Invalid Names by ICZN, n. comb. 1857) n. comb. 1999: Opinion 1030. Sulciconus Bielz, 1869; type species (here anaglyptica (Crosse, 1865) pfluegeri (Petuch, 2003) designated): Conus sulcatus Hwass in Bruguie` re, 1792, n. syn. n. comb. n. comb. The names Asprella and Sulciconus were both published in 1869 arawak (Petuch & R. F. Myers, poremskii (Petuch & and their exact dates of publication are not known in order to es- 2014) n. comb. R. F. Myers, 2014) n. comb. tablish priority. Under ICZN (1999): Art. 24.2, we act here as baccata (G. B. Sowerby III, puncticulata (Hwass in First Revisers and give precedence to the name Asprella over 1877) n. comb. Bruguie` re, 1792) n. comb. Sulciconus.] Downloaded from berschaueri (Petuch & R. F. pusio (Hwass in Bruguie` re, Myers, 2014) n. comb. 1792) n. comb. Included species: damasoi (Cossignani, 2007) roatanensis (Petuch & Sargent, fijisulcatus Moolenbeek, Ro¨ckel helgae Blo¨cher, 1992 n. comb. 2011) n. comb. & Bouchet, 2008 iodostoma Reeve, 1843 ericmonnieri (Petuch & sargenti (Petuch, 2013) gigasulcatus Moolenbeek, rolani Ro¨ckel, 1986 R. F. Myers, 2014) n. comb. n. comb. Ro¨ckel & Bouchet, 2008 sulcatus Hwass in Bruguie` re, http://mollus.oxfordjournals.org/ fluviamaris (Petuch & Sargent, simonei (Petuch & R. F. Myers, grangeri G. B. Sowerby III, 1792 2011) n. comb. 2014) n. comb. 1900 sulcocastaneus Kosuge, 1981 henckesi (Coltro, 2004) n. comb. stearnsii (Conrad, 1869) herndli (Petuch & R. F. Myers, n. comb. 2014) n. comb. tornata (G. B. Sowerby I, Subgenus Atlanticonus Petuch & Sargent, 2012 jaspidea (Gmelin, 1791) 1833) n. comb. n. comb. vanhyningi (Rehder, 1944) Type species: Conus granulatus Linnaeus, 1758; OD lucidus (W. Wood, 1828) n. comb. n. comb. wendrosi (Tenorio & Afonso, Included species: mahogani (Reeve, 1843) 2013) n. comb. cuna Petuch, 1998 granulatus Linnaeus, 1758 by guest on December 3, 2014 n. comb. ximenes (Gray, 1839) glenni Petuch, 1993 ritae Petuch, 1995 mindana (Hwass in n. comb. Bruguie` re, 1792) n. comb. Subgenus Attenuiconus Petuch, 2013 Incertae sedis Type species: Conus attenuatus Reeve, 1844; OD Conasprella delessertii (Rećluz, 1843) n. comb. Included species: attenuatus Reeve, 1844 honkeri Petuch, 1988 Genus Conus Linnaeus, 1758 aureonimbosus Petuch, 1987 poulosi Petuch, 1993 eversoni Petuch, 1987 Subgenus Conus Linnaeus, 1758 Subgenus Austroconus Tucker & Tenorio, 2009 Type species: Conus marmoreus Linnaeus, 1758; SD, Children (1823: 69) Type species: Conus cyanostoma A. Adams, 1855; OD

[Synonyms: Cucullus Ro¨ding, 1798; type species: Conus mar- [Possible senior synonym: Mamiconus Cotton & Godfrey, 1932; moreus Linnaeus, 1758; SD, Winckworth (1945: 139). Coronaxis type species: Conus superstes Hedley, 1911; OD. The identity of C. Swainson, 1840; type species: Conus bandanus Hwass, 1792; M] superstes is uncertain; it is listed as a nomen dubium by Tucker & Tenorio (2013)] Included species: araneosus [Lightfoot], 1786 marmoreus Linnaeus, 1758 Included species: bandanus Hwass in nocturnus [Lightfoot], 1786 clarus E. A. Smith, 1881 sydneyensis G. B. Sowerby III, Bruguie` re, 1792 vidua Reeve, 1843 cyanostoma A. Adams, 1855 1887

Subgenus Afonsoconus Tucker & Tenorio, 2013 Subgenus Bermudaconus Petuch, 2013 Type species: Conus kinoshitai Kuroda, 1956; OD Type species: Conus lightbourni Petuch, 1986; OD Included species: Included species: bruuni Powell, 1958 kinoshitai Kuroda, 1956 lightbourni Petuch, 1986

6 CLASSIFICATION OF CONE SNAILS

Subgenus Brasiliconus Petuch, 2013 [Synonyms: Regiconus Iredale, 1930; type species: Conus Type species: Conus scopulorum Van Mol, Tursch & Kempf, aulicus Linnaeus, 1758; OD; the simultaneously published 1971; OD Darioconus was given precedence over Regiconus by First Reviser Kira (1954: 75). Erythroconus da Motta, 1991; type species: Included species: Conus crocatus Lamarck, 1810; OD] scopulorum Van Mol, Tursch & Kempf, 1971 Included species: aulicus Linnaeus, 1758 leviteni (Tucker, Tenorio & Subgenus Calibanus da Motta, 1991 auratinus da Motta, 1982 Chaney, 2011) n. comb. auricomus Hwass in lohri Kilburn, 1972 Type species: Conus furvus Reeve, 1843; OD Bruguie` re, 1792 madagascariensis G. B. Sowerby behelokensis Lauer, 1989 II, 1858 [Synonym: Thalassiconus Tucker & Tenorio, 2013; type biancae Bozzetti, 2010 magniﬁcus Reeve, 1843 species: Conus thalassiarchus G.B. Sowerby I, 1834, n. syn.; crocatus Lamarck, 1810 natalaurantius (S. G. Veldsman, OD] echo Lauer, 1989 2013) n. comb. Included species: episcopatus da Motta, 1982 omaria Hwass in Bruguie` re, furvus Reeve, 1843 thalassiarchus lamberti Souverbie, 1877 1792 G. B. Sowerby I, 1834 laueri (Monnier & Limpalae¨r, pennaceus Born, 1778 2013) n. comb. thomae Gmelin, 1791

Remarks: The species C. thomae was placed in the subgenus Downloaded from Subgenus Chelyconus Mo¨rch, 1852 Darioconus (following Tucker & Tenorio 2013), even though the Type species: Conus testudinarius Hwass, 1792 [synonym of Conus corresponding clade was not highly supported. ermineus Born, 1778]; SD, Cossmann (1896: 160) Included species: Subgenus Dauciconus Cotton, 1945 ermineus Born, 1778 purpurascens G. B. Sowerby http://mollus.oxfordjournals.org/ I, 1833 Type species: Conus daucus Hwass, 1792; OD [Synonyms: Gradiconus da Motta, 1991; type species: Conus gradatus W. Wood, 1828; OD; n. syn.; Magelliconus da Motta, Subgenus Cylinder Montfort, 1810 1991; type species: Conus magellanicus Hwass in Bruguie` re, 1792; Type species: Conus textile Linnaeus, 1758; OD. Dubois & OD; n. syn.; Purpuriconus da Motta, 1991; type species: Conus car- Bour (2010) revived the name Cylindrus as from Batsch, 1789 dinalis Hwass in Bruguie` re, 1792; OD; n. syn.; Cariboconus and designated Conus textile as its type species. This has the conse- Petuch, 2003; type species: Conus amphiurgus Dall, 1889; OD; quence of destabilizing the pulmonate genus name Cylindrus Conasprelloides Tucker & Tenorio, 2009; type species: Conus Fitzinger, 1833, which is in current use. The ICZN will be peti- cancellatus Hwass in Bruguie` re, 1792; OD; n. syn.; tioned to maintain usage of Cylinder Montfort, 1810, and Poremskiconus Petuch, 2013; type species: Conus ziczac Mu¨hlfeld, by guest on December 3, 2014 Cylindrus Fitzinger, 1833, in their accustomed sense.] 1816; OD; n. syn.; Tuckericonus Petuch 2013; type species: Conus flavescens G. B. Sowerby I, 1834; OD; n. syn.] Included species: abbas Hwass in Bruguie` re, 1792 legatus Lamarck, 1810 Included species: ammiralis Linnaeus, 1758 nodulosus G. B. Sowerby II, abrolhosensis Petuch, 1987 flavescens G. B. Sowerby I, aureus Hwass in Bruguie` re, 1864 alainallaryi Bozzetti & 1834 1792 priscai (Bozzetti, 2012) Monnier, 2009 garciai da Motta, 1982 barbieri G. Raybaudi Massilia, n. comb. amphiurgus Dall, 1889 gibsonsmithorum Petuch, 1986 1995 retifer Menke, 1829 anabathrum Crosse, 1865 glicksteini Petuch, 1987 bengalensis (Okutani, 1968) scottjordani (Poppe, Monnier & arangoi Sarasuá, 1977 goajira Petuch, 1992 canonicus Hwass in Tagaro, 2012) n. comb. aureopunctatus Petuch, 1987 gradatus W. Wood, 1828 Bruguie` re, 1792 tagaroae (Limpalae¨r& bayeri Petuch, 1987 harasewychi Petuch, 1987 dalli Stearns, 1873 Monnier, 2013) n. comb. belizeanus (Petuch & Sargent, havanensis Aguayo & Pe´rez gloriamaris Chemnitz, 1777 telatus Reeve, 1848 2011) n. comb. Farfante, 1947 glorioceanus Poppe & Tagaro, textile Linnaeus, 1758 bessei Petuch, 1992 hennequini Petuch, 1993 2009 victoriae Reeve, 1843 boui da Motta, 1988 hilli Petuch, 1990 brunneobandatus Petuch, 1992 honkerorum (Petuch & Remarks: Tucker & Tenorio (2009) described two ‘body plans’ in brunneofilaris Petuch, 1990 R. F. Myers, 2014) n. comb. their genus Cylinder, citing Conus textile and C. gloriamaris as exam- burryae Clench, 1942 inconstans E. A. Smith, 1877 ples of each group; they may correspond to the two groups dianthus G. B. Sowerby III, jacarusoi Petuch, 1998 found in the molecular phylogeny. Tucker & Tenorio (2009) 1882 jorioi (Petuch, 2013) n. comb. relied on their similar radulae and diets to maintain them in a dispar G. B. Sowerby I, 1833 jucundus G. B. Sowerby III, single genus: we have followed them and place all the species in donnae Petuch, 1998 1887 a single subgenus. Among the sequenced species, C. ammiralis, C. edwardpauli Petuch, 1998 kaiserae (Tenorio, Tucker & canonicus and C. dalli cluster with the type species C. textile; the eleutheraensis (Petuch, 2013) Chaney, 2012) n. comb. other species constitute a separate clade. n. comb. kalafuti da Motta, 1987 ernesti Petuch, 1990 kerstitchi Walls, 1978 explorator Vink, 1990 kevani Petuch, 1987 Subgenus Darioconus Iredale, 1930 flamingo Petuch, 1980 kirkandersi Petuch, 1987 flammeacolor Petuch, 1992 kremerorum Petuch, 1988 Type species: Conus omaria Hwass, 1792; OD kulkulcan Petuch, 1980

7 N. PUILLANDRE ET AL. largilliertii Kiener, 1847 patglicksteinae Petuch, 1987 Subgenus Elisaconus Tucker & Tenorio, 2013 leekremeri Petuch, 1987 paulae Petuch, 1988 Type species: Conus litteratus Linnaeus, 1758; OD levistimpsoni (Tucker, 2013) penchaszadehi Petuch, 1986 n. comb. philippii Kiener, 1847 Included species: lucaya Petuch, 2000 poormani Berry, 1968 litteratus Linnaeus, 1758 maculiferus G. B. Sowerby I, portobeloensis Petuch, 1990 1833 pseudocardinalis Coltro, 2004 magellanicus Hwass in recurvus Broderip, 1833 Subgenus Embrikena Iredale, 1937 Bruguie` re, 1792 regularis G. B. Sowerby I, 1833 Type species: Embrikena pergrandis Iredale, 1937; M magnottei Petuch, 1987 richardbinghami Petuch, 1993 maya (Petuch & Sargent, 2011) rosalindensis Petuch, 1998 Included species: n. comb. rosemaryae Petuch, 1990 darkini Ro¨ckel, Korn & moncuri Filmer, 2005 monilifer Broderip, 1833 sahlbergi da Motta & Harland, Richard, 1993 pergrandis (Iredale, 1937) cancellatus Hwass in 1986 kostini Filmer, Monteiro, tisii T. C. Lan, 1978 Bruguie` re, 1792 scalaris Valenciennes, 1832 Lorenz & Verdasca, 2012 cardinalis Hwass in Bruguie` re, scalarissimus da Motta, 1988 1792 sennottorum Rehder & Abbott, cargilei Coltro, 2004 1951 Subgenus Eremiconus Tucker & Tenorio, 2009 castaneus Kiener, 1848 shaskyi (Tenorio, Tucker & Type species: Conus minnamurra (Garrard, 1961); OD caysalensis L. Raybaudi & Chaney, 2012) n. comb.

Prati, 1994 skoglundae (Tenorio, Tucker & Included species: Downloaded from ceruttii Cargile, 1997 Chaney, 2012) n. comb. albellus Ro¨ckel & Korn, 1990 lizardensis Crosse, 1865 cingulatus Lamarck, 1810 sphacelatus G. B. Sowerby I, colmani Ro¨ckel & Korn, 1990 minnamurra (Garrard, 1961) colombi (Monnier & 1833 limpusi Ro¨ckel & Korn, 1990 Limpalae¨r, 2012) n. comb. stanfieldi Petuch, 1998 colombianus Petuch, 1987 stimpsoni Dall, 1902 Subgenus Eugeniconus da Motta, 1991 coltrorum (Petuch & sunderlandi Petuch, 1987 http://mollus.oxfordjournals.org/ R. F. Myers, 2014) n. comb. theodorei Petuch, 2000 Type species: Conus nobilis Linnaeus, 1758; OD daucus Hwass in Bruguie` re, tristensis Petuch, 1987 1792 velaensis Petuch, 1993 Included species: deynzerorum Petuch, 1995 venezuelanus Petuch, 1987 cordigera G. B. Sowerby II, marchionatus Hinds, 1843 nybakkeni (Tenorio, Tucker & vikingorum Petuch, 1993 1866 nobilis Linnaeus, 1758 Chaney, 2012) n. comb. villepinii P. Fischer & orion Broderip, 1833 Bernardi, 1857 Subgenus Floraconus Iredale, 1930 ortneri Petuch, 1998 virgatus Reeve, 1849 Type species: Conus anemone Lamarck, 1810; OD ostrinus (Tucker & Tenorio, vittatus Hwass in Bruguie` re, 2011) n. comb. 1792 [Synonym: Ketyconus da Motta, 1991; type species: Conus tinia- by guest on December 3, 2014 paraguana Petuch, 1987 xanthicus Dall, 1910 nus Hwass in Bruguie` re, 1792; OD; n. syn.] parascalaris Petuch, 1987 ziczac Mu¨hlfeld, 1816 paschalli Petuch, 1998 zylmanae Petuch, 1998 Included species: anemone Lamarck, 1810 gabelishi da Motta & Subgenus Dendroconus Swainson, 1840 aplustre Reeve, 1843 Ninomiya, 1982 ardisiaceus Kiener, 1850 mozambicus Hwass in Type species: Conus betulinus Linnaeus, 1758; SD, balteatus G. B. Sowerby I, Bruguie` re, 1792 Herrmannsen (1847: 177) 1833 papilliferus G. B. Sowerby I, cocceus Reeve, 1844 1834 [Synonym: Cleobula Iredale, 1930; type species: Conus figuli- compressus G. B. Sowerby tinianus Hwass in Bruguie` re, nus Linnaeus, 1758; OD] II, 1866 1792 Included species: betulinus Linnaeus, 1758 medoci Lorenz, 2004 Subgenus Fraterconus Tucker & Tenorio, 2013 buxeus (Ro¨ding, 1798) suratensis Hwass in Bruguie` re, Type species: Conus distans Hwass in Bruguie` re, 1792; OD figulinus Linnaeus, 1758 1792 glaucus Linnaeus, 1758 Included species: distans Hwass in Bruguie` re, 1792 Remarks: The species C. medoci was placed in the subgenus Dendroconus (following Tucker and Tenorio 2013), even if the corresponding clade was not highly supported Subgenus Gastridium Modeer, 1793 Type species: Conus geographus Linnaeus, 1758; SD, Subgenus Ductoconus da Motta, 1991 Winckworth (1935: 322) Type species: Conus princeps Linnaeus, 1758; OD [Synonyms: Rollus Montfort, 1810; type species: Conus geographus Linnaeus, 1758; OD. Utriculus Schumacher, 1817; type [Synonym: Arubaconus Petuch, 2013; type species: Conus hiero- species: Conus geographus Linnaeus, 1758; M. Tuliparia glyphus Duclos, 1833; OD; n. syn.] Swainson, 1840; type species: Coronaxis nebulosa Swainson, 1840 [synonym of Conus tulipa]; M. Protostrioconus Tucker & Included species: Tenorio, 2009; type species: Conus obscurus G.B. Sowerby I, hieroglyphus Duclos, 1833 princeps Linnaeus, 1758 1833; OD; n. syn.]

8 CLASSIFICATION OF CONE SNAILS

Included species: Included species: cuvieri Crosse, 1858 obscurus G. B. Sowerby I, carcellesi Martins, 1945 patriceae (Petuch & eldredi Morrison, 1955 1833 clerii Reeve, 1844 R. F. Myers, 2014) n. comb. fragilissimus Petuch, 1979 tulipa Linnaeus, 1758 lemniscatus Reeve, 1849 tostesi Petuch, 1986 geographus Linnaeus, 1758 xanthocinctus Petuch, 1986

Subgenus Harmoniconus da Motta, 1991 Subgenus Lautoconus Monterosato, 1923 Type species: Conus musicus Hwass in Bruguie` re, 1792; OD Type species: Conus mediterraneus Hwass in Bruguie` re, 1792 [synonym of Conus ventricosus Gmelin, 1791]; OD Included species: mcbridei Lorenz, 2005 parvatus Walls, 1979 [Synonyms: Africonus Petuch, 1975; type species: Conus cuneo- musicus Hwass in Bruguie` re, paukstisi (Tucker, Tenorio & lus Reeve, 1843; OD; n. syn. Varioconus da Motta, 1991; type 1792 Chaney, 2011) n. comb. species: Conus variegatus Kiener, 1848 OD; n. syn.] nanus G. B. Sowerby I, 1833 sponsalis Hwass in nux Broderip, 1833 Bruguie` re, 1792 Included species: aemulus Reeve, 1844 grahami Ro¨ckel, Cosel & africanus Kiener, 1848 Burnay, 1980 Subgenus Hermes Montfort, 1810 albuquerquei Trova˜o, 1978 guanche Lauer, 1993

Type species: Conus nussatella Linnaeus, 1758; OD alexandrinus Kaicher, 1977 guinaicus Hwass in Downloaded from allaryi Bozzetti, 2008 Bruguie` re, 1792 [Synonym: Theliconus Swainson, 1840; type species: Conus anabelae Rolań&Ro¨ckel, 2001 hybridus Kiener, 1847 nussatella Linnaeus, 1758; OD] anthonyi (Petuch, 1975) infinitus Rolań, 1990 antonioi (Cossignani, 2014) irregularis G. B. Sowerby II, Included species: n. comb. 1858 nussatella Linnaeus, 1758

antoniomonteiroi Rolań, 1990 isabelarum Tenorio & Afonso, http://mollus.oxfordjournals.org/ babaensis Rolań&Ro¨ckel, 2001 2004 Subgenus Kalloconus da Motta, 1991 belairensis Pin & Leung Tack in josephinae Rolań, 1980 Pin, 1989 joserochoi (Cossignani, 2014) Type species: Conus pulcher [Lightfoot], 1786; OD bellulus Rolań, 1990 n. comb. [Synonyms: Genuanoconus Tucker & Tenorio, 2009; type species: bernardinoi (Cossignani, 2014) jourdani da Motta, 1984 Conus genuanus Linnaeus, 1758; OD; n. syn. Trovaoconus n. comb. kersteni Tenorio, Afonso & Tucker & Tenorio, 2009; type species: Conus venulatus Hwass boavistensis Rolań& Rolań, 2008 in Bruguie` re, 1792; OD; n. syn.] Fernandes, 1990 lineopunctatus Kaicher, 1977 Included species: bocagei Trovaõ, 1978 lobitensis Kaicher, 1977 borgesi Trov aõ, 1979 longilineus Ro¨ckel, Rolań&

ateralbus Kiener, 1850 pseudonivifer Monteiro, by guest on December 3, 2014 atlanticoselvagem Afonso & Tenorio & Poppe, 2004 bruguieri Kiener, 1846 Monteiro, 1980 Tenorio, 2004 pulcher [Lightfoot], 1786 bulbus Reeve, 1843 lugubris Reeve, 1849 byssinus (Ro¨ding, 1798) trochulus Reeve, 1844 cabraloi (Cossignani, 2014) luquei Rolań&Trovaõ, 1990 genuanus Linnaeus, 1758 venulatus Hwass in n. comb. maioensis Trovaõ, Rolań& Bruguie` re, 1792 cacao Ferrario, 1983 Fe´lix-Alves, 1990 cagarralensis (Cossignani, 2014) melissae Tenorio, Afonso & n. comb. Rolań, 2008 Subgenus Kellyconus Petuch, 2013 evorai Monteiro, Fernandes mercator Linnaeus, 1758 & Rolań, 1995 messiasi Rolań & Fernandes, Type species: Conus patae Abbott, 1971; OD fantasmalis Rolań, 1990 1990 felitae Rolań, 1990 micropunctatus Rolań&Ro¨ckel, Included species: fernandesi Tenorio, Afonso & 2000 binghamae Petuch, 1987 patae Abbott, 1971 Rolań, 2008 calhetae Rolań, 1990 ignotus Cargile, 1998 fiadeiroi (Tenorio, Afonso, cepasi Trovaõ, 1975 Cunha & Rolań, 2014) chytreus Tryon, 1884 n. comb. claudiae Tenorio & Afonso, Subgenus Klemaeconus Tucker & Tenorio, 2013 filmeri Rolań&Ro¨ckel, 2000 2004 Type species: Rhizoconus klemae Cotton, 1953; OD flavusalbus Rolań&Ro¨ckel, cloveri Walls, 1978 2000 condei (Afonso & Tenorio, [Synonym: Continuconus Tucker & Tenorio, 2013; type species: fontonae Rolań&Trovaõ, 2014) n. comb. Conus plinthis Richard & Moolenbeek, 1988; OD; n. syn.] 1990 crioulus Tenorio & Afonso, franciscoi Rolań&Ro¨ckel, 2004 Included species: 2000 crotchii Reeve, 1849 estivali Moolenbeek & Richard, plinthis Richard & furnae Rolań, 1990 cuneolus Reeve, 1843 1995 Moolenbeek, 1988 fuscoflavus Ro¨ckel, Rolań& curralensis Rolań, 1986 hirasei (Kuroda, 1956) rufimaculosus Macpherson, 1959 Monteiro, 1980 damottai Trovaõ, 1979 klemae (Cotton, 1953) sugimotonis Kuroda, 1928 fuscolineatus G. B. Sowerby III, decoratus Ro¨ckel, Rolań& 1905 Monteiro, 1980 gonsaloi (Afonso & Tenorio, delanoyae Trovaõ, 1979 Subgenus Lamniconus da Motta, 1991 2014) n. comb. denizi (Afonso & Tenorio, Type species: Conus clerii Reeve, 1844; OD 2011) n. comb.

9 N. PUILLANDRE ET AL. derrubado Rolań& santanaensis (Afonso & Tenorio, Included species: Fernandes, 1990 2014) n. comb. bahamensis Vink & Ro¨ckel, spurius Gmelin, 1791 desidiosus A. Adams, 1853 saragasae Rolań, 1986 1995 therriaulti (Petuch, 2013) diegoi (Cossignani, 2014) serranegrae Rolań, 1990 lindae Petuch, 1987 n. comb. n. comb. silviae (Cossignani, 2014) diminutus Trovaõ& Rolań, n. comb. 1986 swinneni (Tenorio, Afonso, Subgenus Lithoconus Mo¨rch, 1852 dorotheae Monnier & Cunha & Rolań, 2014) Limpalae¨r, 2010 n. comb. Type species: Conus millepunctatus Lamarck, 1822 [synonym of echinophilus (Petuch, 1975) tacomae Boyer & Pelorce, 2009 Conus leopardus Ro¨ding, 1798]; SD, Cossmann (1896: 186) miruchae Ro¨ckel, Rolań& taslei Kiener, 1850 Monteiro, 1980 tenuilineatus Rolań&Ro¨ckel, Included species: mordeirae Rolań&Trovaõ, 2001 leopardus Ro¨ding, 1798 1990 teodorae Rolań & Fernandes, naranjus Trovaõ, 1975 1990 navarroi Rolań, 1986 trencarti Nolf & Verstraeten, Subgenus Lividoconus Wils, 1970 negroides Kaicher, 1977 2008 Type species: Conus lividus Hwass in Bruguie` re, 1792; OD nobrei Trovaõ, 1975 trovaoi Rolań&Ro¨ckel, 2000 pineaui Pin & Leung Tack, unifasciatus Kiener, 1850 [Synonym: Calamiconus Tucker & Tenorio, 2009; type species: 1989 variegatus Kiener, 1848 Conus lischkeanus Weinkauff, 1875; OD; n. syn.] pseudocuneolus Ro¨ckel, vayssierei Pallary, 1906 Downloaded from Rolań & Monteiro, 1980 ventricosus Gmelin, 1791 Included species: raulsilvai Rolań, Monteiro & verdensis Trovaõ, 1979 diadema G. B. Sowerby I, kermadecensis Iredale, 1913 Fernandes, 1998 vulcanus Tenorio & Afonso, 1834 lischkeanus Weinkauff, 1875 regonae Rolań&Trovaõ, 2004 escondidai Poppe & Tagaro, lividus Hwass in Bruguie` re, 1990 wandae (Cossignani, 2014) 2005 1792

roeckeli Rolań, 1980 n. comb. eximius Reeve, 1849 muriculatus G. B. Sowerby http://mollus.oxfordjournals.org/ salletae (Cossignani, 2014) xicoi Ro¨ckel, 1987 floridulus A. Adams & I, 1833 n. comb. zebroides Kiener, 1848 Reeve, 1848 quercinus [Lightfoot], 1786 salreiensis Rolań, 1980 zinhoi (Cossignani, 2014) garywilsoni Lorenz & Morrison, sanguinolentus Quoy & n. comb. 2004 Gaimard, 1834

Subgenus Leporiconus Iredale, 1930 Subgenus Monteiroconus da Motta, 1991

Type species: Conus glans Hwass in Bruguie` re, 1792; OD Type species: Conus ambiguus Reeve, 1844; OD

Included species: [Synonym: Gladioconus Tucker & Tenorio, 2009; type species: by guest on December 3, 2014 coffeae Gmelin, 1791 luteus G. B. Sowerby I, 1833 Conus gladiator Broderip, 1833; OD; n. syn.] glans Hwass in Bruguie` re, tenuistriatus G. B. Sowerby 1792 II, 1858 Included species: granum Ro¨ckel & Fischo¨der, ambiguus Reeve, 1844 mus Hwass in Bruguie` re, 1792 1985 bellocqae van Rossum, 1996 tabidus Reeve, 1844 gladiator Broderip, 1833

Subgenus Leptoconus Swainson 1840 Subgenus Papyriconus Tucker & Tenorio, 2013

Type species: Conus amadis Gmelin, 1791; SD, Herrmannsen Type species: Conus sculletti Marsh, 1962; OD (1847: 584) Included species: [Synonym: Nataliconus Tucker & Tenorio, 2009; type species: sculletti Marsh, 1962 Conus natalis G. B. Sowerby II, 1858; OD; n. syn.]

Included species: Subgenus Phasmoconus Mo¨rch, 1852 amadis Gmelin, 1791 milneedwardsi Jousseaume, 1894 hamanni Fainzilber & Mienis, natalis G. B. Sowerby II, Type species: Conus radiatus Gmelin, 1791; SD, Cotton (1945: 250) 1986 1858 immelmani Korn, 1998 royaikeni (S. G. Veldsman, [Synonyms: Fulgiconus da Motta, 1991; type species: Conus kawamurai Habe, 1962 2010) n. comb. moluccensis Ku¨ster, 1838; OD; n. syn.; Graphiconus da Motta, locumtenens Blumenbach, 1991; type species: Conus inscriptus Reeve, 1843; OD; n. syn.; 1791 Thoraconus da Motta, 1991; type species: Conus exiguus Lamarck, 1810; OD; n. syn. Nimboconus Tucker & Tenorio, 2013;type species: Conus nimbosus Hwass in Bruguie` re, 1792; OD; n. syn.] Subgenus Lindaconus Petuch, 2002 Included species: Type species: Conus lindae Petuch, 1987; OD alabaster Reeve, 1849 andamanensis E. A. Smith, alexandrei (Limpalae¨r& 1879 [Synonym: Spuriconus Petuch, 2003; type species: Conus Monnier, 2012) n. comb. angioiorum Ro¨ckel & spurius Gmelin, 1791; OD] amplus Ro¨ckel & Korn, 1992 Moolenbeek, 1992

10 CLASSIFICATION OF CONE SNAILS armadillo Shikama, 1971 conspersus Reeve, 1844 consors G. B. Sowerby I, leobottonii Lorenz, 2006 asiaticus da Motta, 1985 dampierensis Coomans & 1833 magus Linnaeus, 1758 australis Holten, 1802 Filmer, 1985 fischoederi Ro¨ckel & da Motta, monachus Linnaeus, 1758 balabacensis Filmer, 2012 daphne Boivin, 1864 1983 morrisoni G. Raybaudi blanfordianus Crosse, 1867 erythraeensis Reeve, 1843 floccatus G. B. Sowerby I, Massilia, 1991 broderipii Reeve, 1844 evansi Bondarev, 2001 1841 nigromaculatus Ro¨ckel & cavailloni Fenaux, 1942 exiguus Lamarck, 1810 fulmen Reeve, 1843 Moolenbeek, 1992 cebuensis Wils, 1990 flavus Ro¨ckel, 1985 gauguini Richard & Salvat, nigropunctatus G. B. Sowerby II, ciderryi da Motta, 1985 gilvus Reeve, 1849 1973 1858 cinereus Hwass in Bruguie` re, giorossii Bozzetti, 2005 stercusmuscarum robini (Limpalae¨r & Monnier, 1792 goudeyi (Monnier & Limpalae¨r, Linnaeus, 1758 2012) n. comb. collisus Reeve, 1849 2012) n. comb. striatus Linnaeus, 1758 rouxi (Monnier, Limpalae¨r& laterculatus G. B. Sowerby habui Lan, 2002 gubernator Hwass in Robin, 2013) n. comb. II, 1870 inscriptus Reeve, 1843 Bruguie` re, 1792 striolatus Kiener, 1848 leobrerai da Motta & Martin, janus Hwass in Bruguie` re, koukae (Monnier, Limpalae¨r& 1982 1792 Robin, 2013) n. comb. lienardi Bernardi & Crosse, jickelii Weinkauff, 1873 1861 kiicumulus Azuma, 1982 lynceus G. B. Sowerby II, kuroharai (Habe, 1965) Subgenus Plicaustraconus Moolenbeek, 2008 1858 ranonganus da Motta, 1978

madecassinus (Bozzetti, richeri Richard & Type species: Conus advertex (Garrard, 1961); OD Downloaded from 2012) n. comb. Moolenbeek, 1988 marielae Rehder & Wilson, salzmanni G. Raybaudi Included species: 1975 Massilia & Rola´n, 1997 adami Wils, 1988 felix Fenzan, 2012 martinianus Reeve, 1844 santinii (Monnier & advertex (Garrard, 1961) lozeti Richard, 1980 moluccensis Ku¨ster, 1838 Limpalae¨r, 2014) n. comb. angasi Tryon, 1884 trigonus Reeve, 1848 baeri Ro¨ckel & Korn, 1992 visagenus Kilburn, 1974 moolenbeeki Filmer, 2011 sartii Korn, Niederho¨fer & http://mollus.oxfordjournals.org/ mucronatus Reeve, 1843 Blo¨cher, 2002 bonﬁgliolii (Bozzetti, 2010) wallangra (Garrard, 1961) mulderi Fulton, 1936 scalptus Reeve, 1843 n. comb. neptunus Reeve, 1843 sculpturatus Ro¨ckel & da Motta, niederhoeferi (Monnier, 1986 Limpalae¨r & Lorenz, 2012) sertacinctus Ro¨ckel, 1986 Subgenus Pseudolilliconus Tucker & Tenorio, 2009 n. comb. sogodensis (Poppe, Monnier & nimbosus Hwass in Tagaro, 2012) n. comb. Type species: Conus boschorum Moolenbeek & Coomans, 1993; OD Bruguie` re, 1792 solomonensis Delsaerdt, 1992 ochroleucus Gmelin, 1791 spectrum Linnaeus, 1758 Included species: oishii (Shikama, 1977) stramineus Lamarck, 1810 boschorum Moolenbeek & scalarispira (Bozzetti, 2012) parius Reeve, 1844 straturatus G. B. Sowerby II, Coomans, 1993 n. comb. by guest on December 3, 2014 petergabrieli Lorenz, 2006 1865 korni G. Raybaudi Massilia, traillii A. Adams, 1855 pica A. Adams & Reeve, 1848 sutanorcum Moolenbeek, 1993 visseri Delsaerdt, 1990 pretiosus G. Nevill & H. Nevill, Ro¨ckel & Bouchet, 2008 kuiperi Moolenbeek, 2006 wallacei (Lorenz & Morrison, 1874 vappereaui Monteiro, 2009 levis (Bozzetti, 2012) n. comb. 2004) primus Ro¨ckel & Korn, 1990 wittigi Walls, 1977 proximus G. B. Sowerby II, yemenensis Bondarev, 1997 1860 zandbergeni Filmer & Subgenus Pseudonoduloconus Tucker & Tenorio, 2009 quiquandoni Lorenz & Barbier, Moolenbeek, 2010 2008 zapatosensis Ro¨ckel, 1987 Type species: Conus carnalis G. B. Sowerby III, 1879; OD radiatus Gmelin, 1791 zebra Lamarck, 1810 Included species: carnalis G. B. Sowerby III, 1879 Subgenus Pionoconus Mo¨rch, 1852 Type species: Conus magus Linnaeus, 1758; SD, Cossmann Subgenus Pseudopterygia Tucker & Tenorio, 2013 (1896: 160) Type species: Conus terryni Tenorio & Poppe, 2004; OD [Synonyms: Heroconus da Motta, 1991; type species: Conus poehlia- nus G. B. Sowerby III, 1887 [synonym of Conus consors Included species: G. B. Sowerby I, 1833]; OD. Strioconus Thiele, 1929; type species: terryni Tenorio & Poppe, 2004 Conus striatus Linnaeus, 1758; OD. Socioconus da Motta, 1991; type species: Conus consors G. B. Sowerby I, 1833; OD] Subgenus Puncticulis Swainson, 1840 Included species: achatinus Gmelin, 1791 barthelemyi Bernardi, 1861 Type species: Conus arenatus Hwass in Bruguie` re, 1792; M. arafurensis (Monnier, boutetorum Richard & Rabiller, Limpalae¨r & Robin, 2013) 2013 Included species: n. comb. catus Hwass in Bruguie` re, arenatus Hwass in Bruguie` re, vautieri Kiener, 1847 atimovatae (Bozzetti, 2012) 1792 1792 caracteristicus Fischer von n. comb. circumcisus Born, 1778 pulicarius Hwass in Waldheim, 1807 aurisiacus Linnaeus, 1758 Bruguie` re, 1792 zeylanicus Gmelin, 1791

11 N. PUILLANDRE ET AL.

Subgenus Pyruconus Olsson, 1967 Subgenus Splinoconus da Motta, 1991

Type species: Conus patricius Hinds, 1843; OD Type species: Conus biliosus Ro¨ding, 1798; OD Included species: [Synonyms: Kioconus da Motta, 1991; type species: Conus fergusoni G. B. Sowerby II, patricius Hinds, 1843 recluzianus Bernardi, 1853; OD; n. syn. Ongoconus da Motta, 1873 1991; type species: Conus voluminalis Reeve, 1843; OD; Precedence of Splinoconus (established at the rank of genus) over Remarks: Even though the two species are not in the same group in simultaneously established Kioconus and Ongoconus (established the molecular phylogeny, and to avoid the creation of a new sub- at the rank of subgenus) determined by ICZN (1999) Art. 24.1. generic name, we have followed Tucker & Tenorio (2009),who Stellaconus Tucker & Tenorio, 2009; type species: Conus malacanus placed C. fergusoni and C. patricius in the same genus, Pyruconus Hwass in Bruguie` re, 1792; OD; n. syn. Isoconus Tucker & (here ranked as subgenus), based on radula and shell characters. Tenorio, 2013; type species: Conus corallinus Kiener, 1847; OD; n. syn. Nitidoconus Tucker & Tenorio, 2013; type species: Subgenus Quasiconus Tucker & Tenorio, 2009 Conus boeticus Reeve, 1844; OD; n. syn.] Type species: Conus melvilli G. B. Sowerby III, 1879; OD Included species: alconnelli da Motta, 1986 typhon Kilburn, 1975 Included species: bayani Jousseaume, 1872 martensi E. A. Smith, 1884 melvilli G. B. Sowerby III, tuticorinensis Ro¨ckel & Korn, biliosus (Ro¨ding, 1798) nielsenae Marsh, 1962 1879 1990 boeticus Reeve, 1844 nucleus Reeve, 1848 bondarevi Ro¨ckel & olgiatii Bozzetti, 2007 Downloaded from G. Raybaudi Massilia, 1992 papuensis Coomans & Subgenus Rhizoconus Mo¨rch, 1852 caillaudii Kiener, 1846 Moolenbeek, 1982 Type species: Conus miles Linnaeus, 1758; SD, Cossmann capitanellus Fulton, 1938 pauperculus G. B. Sowerby I, (1896: 162) capreolus Ro¨ckel, 1985 1834 corallinus Kiener, 1847 purissimus Filmer, 2011 Included species: danilai Ro¨ckel & Korn, 1990 queenslandis da Motta, 1984 http://mollus.oxfordjournals.org/ anosyensis Bozzetti, 2008 peli Moolenbeek, 1996 dayriti Ro¨ckel & da Motta, recluzianus Bernardi, 1853 capitaneus Linnaeus, 1758 pertusus Hwass in Bruguie` re, 1983 reductaspiralis Walls, 1979 cumingii Reeve, 1848 1792 empressae Lorenz, 2001 richardsae Ro¨ckel & Korn, 1992 hyaena Hwass in Bruguie` re, rattus Hwass in Bruguie` re, fumigatus Hwass in roseorapum G. Raybaudi & 1792 1792 Bruguie` re, 1792 da Motta, 1990 miles Linnaeus, 1758 rawaiensis da Motta, 1978 gilberti (Bozzetti, 2012) n. comb ruthae (Monnier & Limpalae¨r, mustelinus Hwass in tirardi Ro¨ckel & Moolenbeek, gloriakiiensis Kuroda & Itoˆ, 2013) n. comb. Bruguie` re, 1792 1996 1961 sazanka Shikama, 1970 namocanus Hwass in vexillum Gmelin, 1791 hamamotoi Yoshiba & shikamai Coomans,

Bruguie` re, 1792 wilsi Delsaerdt, 1998 Koyama, 1984 Moolenbeek & Wils, 1985 by guest on December 3, 2014 lenavati da Motta & Ro¨ckel, sukhadwalai Ro¨ckel & da 1982 Motta, 1983 Subgenus Rubroconus Tucker & Tenorio, 2013 malacanus Hwass in Bruguie` re, urashimanus Kuroda & Itoˆ, 1792 1961 Type species: Conus coccineus Gmelin, 1791; OD thevenardensis da Motta, 1987 vanvilstereni (Moolenbeek & tribblei Walls, 1977 Zandbergen, 2013) n. comb. Included species: troendlei Moolenbeek, viola Cernohorsky, 1977 coccineus Gmelin, 1791 Zandbergen & Bouchet, 2008 voluminalis Reeve, 1843

Subgenus Sandericonus Petuch, 2013 Type species: Conus sanderi Wils & Moolenbeek, 1979; OD Subgenus Stephanoconus Mo¨rch, 1852 Type species: Conus leucostictus Gmelin, 1791 [¼ Conus regius Included species: Gmelin, 1791]; SD, Wenz (1943: 1469) carioca Petuch, 1986 ednae (Petuch, 2013) n. comb. [Synonyms: Rhombus Montfort, 1810 (invalid; junior homonym sanderi Wils & Moolenbeek, 1979 of Rhombus Walbaum, 1792); type species: Conus imperialis Linnaeus, 1758; OD. Cornutoconus Suzuki, 1972; type species: Subgenus Sciteconus da Motta, 1991 Cornutoconus lamellatus Suzuki, 1972 (¼ Conus chiangi (Azuma, 1972)); OD. Taranteconus Azuma, 1972; type species: Conus Type species: Conus algoensis G. B. Sowerby I, 1834; OD chiangi (Azuma, 1972); OD; n. syn. Protoconus da Motta, 1991 (invalid; junior homonym of Protoconus Yu, 1979 and Protoconus Included species: Stinchcomb, 1986); type species: Conus cedonulli Linnaeus, algoensis G. B. Sowerby I, 1834 gradatulus Weinkauff, 1875 1767; OD. Seminoleconus Petuch, 2003; type species: †Conus viole- bairstowi G. B. Sowerby III, infrenatus Reeve, 1848 tae Petuch 1988 (Pliocene); OD. Rhombiconus Tucker & Tenorio, 1889 pictus Reeve, 1843 2009; type species: Conus imperialis Linnaeus, 1758; OD; brianhayesi Korn, 2001 n. syn. Tenorioconus Petuch & Drohlshagen, 2011; type species: Conus cedonulli Linnaeus, 1767; OD; n. syn.]

12 CLASSIFICATION OF CONE SNAILS

Included species: Tucker & Tenorio, 2013; type species: Conus mitratus Hwass archon Broderip, 1833 granarius Kiener, 1847 in Bruguie` re, 1792; OD; n. syn.] aurantius Hwass in Bruguie` re, imperialis Linnaeus, 1758 1792 julieandreae Cargile, 1995 Included species: bartschi G. D. Hanna & mappa [Lightfoot], 1786 Strong, 1949 polongimarumai Kosuge, 1980 acutangulus Lamarck, 1810 gratacapii Pilsbry, 1904 brunneus Wood, 1828 pseudaurantius Vink & Cosel, andremenezi Olivera & luciae Moolenbeek, 1986 cedonulli Linnaeus, 1767 1985 Biggs, 2010 milesi E. A. Smith, 1887 chiangi (Azuma, 1972) pseudimperialis Moolenbeek, beatrix Tenorio, Poppe & miniexcelsus Olivera & curassaviensis Hwass in Zandbergen & Bouchet, 2008 Tagaro, 2007 Biggs, 2010 Bruguie` re, 1792 regius Gmelin, 1791 cylindraceus Broderip & mitratus Hwass in Bruguie` re, dominicanus Hwass in sanguineus Kiener, 1850 G. B. Sowerby I, 1830 1792 Bruguie` re, 1792 solidus Gmelin, 1791 excelsus G. B. Sowerby III, praecellens A. Adams, 1855 duffyi Petuch, 1992 zonatus Hwass in Bruguie` re, 1908 rizali Olivera & Biggs, 2010 harlandi Petuch, 1987 1792 gondwanensis Ro¨ckel & stupa (Kuroda, 1956) Moolenbeek, 1995 stupella (Kuroda, 1956) Subgenus Strategoconus da Motta, 1991 Type species: Conus generalis Linnaeus, 1767; OD Subgenus Virgiconus Cotton, 1945 [Synonyms: Vituliconus da Motta, 1991; type species: Conus vituli- Type species: Conus virgo Linnaeus, 1758; OD nus Hwass in Bruguie` re, 1792 (¼ Conus planorbis Born, 1778); Downloaded from OD; n. syn. Rolaniconus Tucker & Tenorio, 2009; type species: [Synonym: Pseudohermes Tucker & Tenorio 2013; type species: Conus varius Linnaeus, 1758; OD; n. syn.] Conus austroviola Ro¨ckel & Korn, 1992; OD; n. syn.] Included species: Included species: athenae Filmer, 2011 circumactus Iredale, 1929 artoptus G. B. Sowerby I, 1833 kintoki Habe & Kosuge, 1970 augur [Lightfoot], 1786 dedonderi (Goethaels & austroviola Ro¨ckel & Korn, moreleti Crosse, 1858 http://mollus.oxfordjournals.org/ axelrodi Walls, 1978 D. Monsecour, 2013) n. comb. 1992 spiceri Bartsch & Rehder, 1943 buniatus (Bozzetti, 2013) ferrugineus Hwass in berdulinus Veillard, 1972 terebra Born, 1778 n. comb. Bruguie` re, 1792 coelinae Crosse, 1858 tethys (Petuch & Sargent, 2011) generalis Linnaeus, 1767 monile Hwass in Bruguie` re, emaciatus Reeve, 1849 n. comb. hanshassi (Lorenz & Barbier, 1792 ﬂavidus Lamarck, 1810 violaceus Gmelin, 1791 2012) n. comb. planorbis Born, 1778 frigidus Reeve, 1848 virgo Linnaeus, 1758 hivanus Moolenbeek, splendidulus G. B. Sowerby I, Zandbergen & Bouchet, 1833 2008 suduirauti Raybaudi Massilia, Subgenus Virroconus Iredale, 1930 litoglyphus Hwass in 2004

Bruguie` re, 1792 striatellus Link, 1807 Type species: Conus ebraeus Linnaeus, 1758; OD by guest on December 3, 2014 maldivus Hwass in Bruguie` re, swainsoni Estival & Cosel, 1986 [Synonyms: Dyraspis Iredale, 1949; type species: Conus dor- 1792 varius Linnaeus, 1758 reensis Pe´ron, 1807; OD, n. syn. Miliariconus Tucker & Tenorio, 2009; type species: Conus miliaris Hwass in Subgenus Tesselliconus da Motta, 1991 Bruguie` re, 1792; OD, n. syn.] Type species: Conus tessulatus Born, 1778; OD Included species: Included species: abbreviatus Reeve, 1843 lecourtorum (Lorenz, 2011) eburneus Hwass in Bruguie` re, sandwichensis Walls, 1978 aristophanes G. B. Sowerby n. comb. 1792 suturatus Reeve, 1844 II, 1857 miliaris Hwass in Bruguie` re, edaphus Dall, 1910 tessulatus Born, 1778 chaldaeus (Ro¨ding, 1798) 1792 coronatus Gmelin, 1791 taeniatus Hwass in Bruguie` re, Subgenus Textilia Swainson, 1840 dorreensis Pe´ron, 1807 1792 ebraeus Linnaeus, 1758 tiaratus G. B. Sowerby I, Type species: Conus bullatus Linnaeus, 1758; SD, Cotton encaustus Kiener, 1845 1833 (1945: 261) judaeus Bergh, 1895 Included species: Incertae Sedis adamsonii Broderip, 1836 lucasi (Bozzetti, 2010) n. comb. bullatus Linnaeus, 1758 solangeae Bozzetti, 2004 Genus Kenyonia Brazier, 1896 cervus Lamarck, 1822 timorensis Hwass in Bruguie` re, Type species: Kenyonia pulcherrima Brazier, 1896; M chiapponorum Lorenz, 2004 1792 dusaveli (H. Adams, 1872) vicweei Old, 1973 Included species: julii Lienard, 1870 pulcherrima Brazier, 1896 Subgenus Turriconus Shikama & Habe, 1968 ACKNOWLEDGEMENTS Type species: Turriconus nakayasui Shikama & Habe, 1968 [synonym of Conus excelsus G. B. Sowerby III, 1908]; OD The authors thank Manuel J. Tenorio for helpful comments. Some of the specimens analysed by BMO for this work were [Synonyms: Kurodaconus Shikama & Habe, 1968; type species: obtained using grant support provided by the National Institutes Embrikena stupa Kuroda, 1956; OD; n. syn. Mitraconus of Health (GM48677 and ICBG grant 1U01TW008163). This

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14 CLASSIFICATION OF CONE SNAILS

APPENDIX Appendix Table 1. Continued Species Genus (Subgenus)

Appendix Table 1. List of the valid Recent species of cones and their armiger (Crosse, 1858) Conasprella (Dalliconus) current classiﬁcation. articulata (G.B. Sowerby II, 1873) Conasprella (Conasprella) Species Genus (Subgenus) artoptus G. B. Sowerby I, 1833 Conus (Virgiconus) asiaticus da Motta, 1985 Conus (Phasmoconus) abbas Hwass in Bruguie`re, 1792 Conus (Cylinder) ateralbus Kiener, 1850 Conus (Kalloconus) abbreviates Reeve, 1843 Conus (Virroconus) athenae Filmer, 2011 Conus (Strategoconus) abrolhosensis Petuch, 1987 Conus (Dauciconus) atimovatae (Bozzetti, 2012) Conus (Pionoconus) achatinus Gmelin, 1791 Conus (Pionoconus) atlanticoselvagem Afonso & Tenorio, 2004 Conus (Kalloconus) aculeiformis (Reeve, 1844) Conasprella (Fusiconus) attenuatus Reeve, 1844 Conus (Attenuiconus) acutangulus Lamarck, 1810 Conus (Turriconus) augur [Lightfoot], 1786 Conus (Strategoconus) adami Wils, 1988 Conus (Plicaustraconus) aulicus Linnaeus, 1758 Conus (Darioconus) adamsonii Broderip, 1836 Conus (Textilia) aurantius Hwass in Bruguie`re, 1792 Conus (Stephanoconus) advertex (Garrard, 1961) Conus (Plicaustraconus) auratinus da Motta, 1982 Conus (Darioconus) aemulus Reeve, 1844 Conus (Lautoconus) aureonimbosus Petuch, 1987 Conus (Attenuiconus) africanus Kiener, 1848 Conus (Lautoconus) aureopunctatus Petuch, 1987 Conus (Dauciconus) alabaster Reeve, 1849 Conus (Phasmoconus) aureus Hwass in Bruguie`re, 1792 Conus (Cylinder) alainallaryi Bozzetti & Monnier, 2009 Conus (Dauciconus) Downloaded from auricomus Hwass in Bruguie`re, 1792 Conus (Darioconus) albellus Ro¨ckel & Korn, 1990 Conus (Eremiconus) aurisiacus Linnaeus, 1758 Conus (Pionoconus) albuquerquei Trova˜o, 1978 Conus (Lautoconus) australis Holten, 1802 Conus (Phasmoconus) alconnelli da Motta, 1986 Conus (Splinoconus) austroviola Ro¨ckel & Korn, 1992 Conus (Virgiconus) alexandrei (Limpalae¨r & Monnier, 2012) Conus (Phasmoconus) axelrodi Walls, 1978 Conus (Strategoconus)

alexandremonteiroi (Cossignani, 2014) Conasprella (Ximeniconus) http://mollus.oxfordjournals.org/ babaensis Rolań&Ro¨ckel, 2001 Conus (Lautoconus) alexandrinus Kaicher, 1977 Conus (Lautoconus) baccata (G. B. Sowerby III, 1877) Conasprella (Ximeniconus) algoensis G. B. Sowerby I, 1834 Conus (Sciteconus) baeri Ro¨ckel & Korn, 1992 Conus (Plicaustraconus) alisi (Moolenbeek, Ro¨ckel & Richard, 1995) Conasprella bahamensis Vink & Ro¨ckel, 1995 Conus (Lindaconus) (Boucheticonus) baileyi (Ro¨ckel & da Motta, 1979) Conasprella (Conasprella) allamandi (Petuch, 2013) Conasprella (Ximeniconus) bairstowi G. B. Sowerby III, 1889 Conus (Sciteconus) allaryi Bozzetti, 2008 Conus (Lautoconus) bajanensis (Nowell-Usticke, 1968) Conasprella (Dalliconus) amadis Gmelin, 1791 Conus (Leptoconus) balabacensis Filmer, 2012 Conus (Phasmoconus) ambiguus Reeve, 1844 Conus (Monteiroconus) balteatus G. B. Sowerby I, 1833 Conus (Floraconus) ammiralis Linnaeus, 1758 Conus (Cylinder) bandanus Hwass in Bruguie`re, 1792 Conus (Conus) amphiurgus Dall, 1889 Conus (Dauciconus) by guest on December 3, 2014 barbieri G. Raybaudi Massilia, 1995 Conus (Cylinder) amplus Ro¨ckel & Korn, 1992 Conus (Phasmoconus) barthelemyi Bernardi, 1861 Conus (Pionoconus) anabathrum Crosse, 1865 Conus (Dauciconus) bartschi G. D. Hanna & Strong, 1949 Conus (Stephanoconus) anabelae Rolań&Ro¨ckel, 2001 Conus (Lautoconus) bayani Jousseaume, 1872 Conus (Splinoconus) anaglyptica (Crosse, 1865) Conasprella (Ximeniconus) bayeri Petuch, 1987 Conus (Dauciconus) andamanensis E. A. Smith, 1879 Conus (Phasmoconus) beatrix Tenorio, Poppe & Tagaro, 2007 Conus (Turriconus) andremenezi Olivera & Biggs, 2010 Conus (Turriconus) behelokensis Lauer, 1989 Conus (Darioconus) anemone Lamarck, 1810 Conus (Floraconus) belairensis Pin & Leung Tack in Pin, 1989 Conus (Lautoconus) angasi Tryon, 1884 Conus (Plicaustraconus) belizeanus (Petuch & Sargent, 2011) Conus (Dauciconus) angioiorum Ro¨ckel & Moolenbeek, 1992 Conus (Phasmoconus) bellocqae van Rossum, 1996 Conus (Monteiroconus) anosyensis Bozzetti, 2008 Conus (Rhizoconus) bellulus Rolań, 1990 Conus (Lautoconus) anthonyi (Petuch, 1975) Conus (Lautoconus) bengalensis (Okutani, 1968) Conus (Cylinder) antonioi (Cossignani, 2014) Conus (Lautoconus) berdulinus Veillard, 1972 Conus (Virgiconus) antoniomonteiroi Rolań, 1990 Conus (Lautoconus) bernardinoi (Cossignani, 2014) Conus (Lautoconus) aphrodite (Petuch, 1979) Conasprella (Conasprella) berschaueri (Petuch & R. F. Myers, 2014) Conasprella (Ximeniconus) aplustre Reeve, 1843 Conus (Floraconus) bessei Petuch, 1992 Conus (Dauciconus) arafurensis (Monnier, Limpalae¨r & Robin, 2013) Conus (Pionoconus) betulinus Linnaeus, 1758 Conus (Dendroconus) araneosus [Lightfoot], 1786 Conus (Conus) biancae Bozzetti, 2010 Conus (Darioconus) arangoi Sarasuá, 1977 Conus (Dauciconus) biliosus (Ro¨ding, 1798) Conus (Splinoconus) arawak (Petuch & R. F. Myers, 2014) Conasprella (Ximeniconus) binghamae Petuch, 1987 Conus (Kellyconus) archon Broderip, 1833 Conus (Stephanoconus) biraghii (G. Raybaudi Massilia, 1992) Conasprella (Lilliconus) arcuata (Broderip & G. B. Sowerby I, 1829) Conasprella (Kohniconus) blanfordianus Crosse, 1867 Conus (Phasmoconus) ardisiaceus Kiener, 1850 Conus (Floraconus) boavistensis Rolań & Fernandes, 1990 Conus (Lautoconus) arenatus Hwass in Bruguie`re, 1792 Conus (Puncticulis) bocagei Trovaõ, 1978 Conus (Lautoconus) aristophanes G. B. Sowerby II, 1857 Conus (Virroconus) boeticus Reeve, 1844 Conus (Splinoconus) armadillo Shikama, 1971 Conus (Phasmoconus) boholensis (Petuch, 1979) Conasprella (Conasprella)

Continued Continued

15 N. PUILLANDRE ET AL.

Appendix Table 1. Continued Appendix Table 1. Continued

Species Genus (Subgenus) Species Genus (Subgenus) bondarevi Ro¨ckel & G. Raybaudi Massilia, 1992 Conus (Splinoconus) cingulatus Lamarck, 1810 Conus (Dauciconus) bonfigliolii (Bozzetti, 2010) Conus (Plicaustraconus) circumactus Iredale, 1929 Conus (Strategoconus) borgesi Trovaõ, 1979 Conus (Lautoconus) circumcisus Born, 1778 Conus (Pionoconus) boschorum Moolenbeek & Coomans, 1993 Conus (Pseudolilliconus) clarus E. A. Smith, 1881 Conus (Mamiconus) boucheti (Richard, 1983) Conasprella (Conasprella) claudiae Tenorio & Afonso, 2004 Conus (Lautoconus) boui da Motta, 1988 Conus (Dauciconus) clerii Reeve, 1844 Conus (Lamniconus) boutetorum Richard & Rabiller, 2013 Conus (Pionoconus) cloveri Walls, 1978 Conus (Lautoconus) bozzettii (Lauer, 1991) Conasprella (Conasprella) cocceus Reeve, 1844 Conus (Floraconus) brianhayesi Korn, 2001 Conus (Sciteconus) coccineus Gmelin, 1791 Conus (Rubroconus) broderipii Reeve, 1844 Conus (Phasmoconus) coelinae Crosse, 1858 Conus (Virgiconus) bruguieri Kiener, 1846 Conus (Lautoconus) coffeae Gmelin, 1791 Conus (Leporiconus) brunneobandatus Petuch, 1992 Conus (Dauciconus) collisus Reeve, 1849 Conus (Phasmoconus) brunneofilaris Petuch, 1990 Conus (Dauciconus) colmani Ro¨ckel & Korn, 1990 Conus (Eremiconus) brunneus Wood, 1828 Conus (Stephanoconus) colombi (Monnier & Limpalae¨r, 2012) Conus (Dauciconus) bruuni Powell, 1958 Conus (Afonsoconus) colombianus Petuch, 1987 Conus (Dauciconus)

bulbus Reeve, 1843 Conus (Lautoconus) coltrorum (Petuch & R. F. Myers, 2014) Conus (Dauciconus) Downloaded from bullatus Linnaeus, 1758 Conus (Textilia) comatosa (Pilsbry, 1904) Conasprella (Fusiconus) buniatus (Bozzetti, 2013) Conus (Strategoconus) compressus G. B. Sowerby II, 1866 Conus (Floraconus) burryae Clench, 1942 Conus (Dauciconus) condei (Afonso & Tenorio, 2014) Conus (Lautoconus) buxeus (Ro¨ding, 1798) Conus (Dendroconus) consors G. B. Sowerby I, 1833 Conus (Pionoconus) byssinus (Ro¨ding, 1798) Conus (Kalloconus) conspersus Reeve, 1844 Conus (Phasmoconus) cabraloi (Cossignani, 2014) Conus (Lautoconus) corallinus Kiener, 1847 Conus (Splinoconus) http://mollus.oxfordjournals.org/ cacao Ferrario, 1983 Conus (Lautoconus) cordigera G. B. Sowerby II, 1866 Conus (Eugeniconus) cagarralensis (Cossignani, 2014) Conus (Lautoconus) coriolisi (Ro¨ckel, Richard & Moolenbeek, 1995) Conasprella (Fusiconus) caillaudii Kiener, 1846 Conus (Splinoconus) coromandelica (E. A. Smith, 1894) Conasprella cakobaui (Moolenbeek, Ro¨ckel & Bouchet, Profundiconus (Pseudoconorbis) 2008) coronatus Gmelin, 1791 Conus (Virroconus) calhetae Rolań, 1990 Conus (Lautoconus) crioulus Tenorio & Afonso, 2004 Conus (Lautoconus) californicus (Reeve, 1844) Californiconus crocatus Lamarck, 1810 Conus (Darioconus) cancellatus Hwass in Bruguie`re, 1792 Conus (Dauciconus) crotchii Reeve, 1849 Conus (Lautoconus) ` canonicus Hwass in Bruguiere, 1792 Conus (Cylinder) cumingii Reeve, 1848 Conus (Rhizoconus) by guest on December 3, 2014 capitanellus Fulton, 1938 Conus (Splinoconus) cuna Petuch, 1998 Conus (Atlanticonus) capitaneus Linnaeus, 1758 Conus (Rhizoconus) cuneolus Reeve, 1843 Conus (Lautoconus) capreolus Ro¨ckel, 1985 Conus (Splinoconus) curassaviensis Hwass in Bruguie`re, 1792 Conus (Stephanoconus) caracteristicus Fischer von Waldheim, 1807 Conus (Puncticulis) curralensis Rolań, 1986 Conus (Lautoconus) carcellesi Martins, 1945 Conus (Lamniconus) cuvieri Crosse, 1858 Conus (Gastridium) cardinalis Hwass in Bruguie`re, 1792 Conus (Dauciconus) cyanostoma A. Adams, 1855 Conus (Mamiconus) cargilei Coltro, 2004 Conus (Dauciconus) cylindraceus Broderip & G. B. Sowerby I, 1830 Conus (Turriconus) carioca Petuch, 1986 Conus (Sandericonus) dalli Stearns, 1873 Conus (Cylinder) carnalis G. B. Sowerby III, 1879 Conus damasoi (Cossignani, 2007) Conasprella (Ximeniconus) (Pseudonoduloconus) damottai Trovaõ, 1979 Conus (Lautoconus) castaneus Kiener, 1848 Conus (Dauciconus) dampierensis Coomans & Filmer, 1985 Conus (Phasmoconus) catus Hwass in Bruguie`re, 1792 Conus (Pionoconus) danilai Ro¨ckel & Korn, 1990 Conus (Splinoconus) cavailloni Fenaux, 1942 Conus (Phasmoconus) daphne Boivin, 1864 Conus (Phasmoconus) caysalensis L. Raybaudi & Prati, 1994 Conus (Dauciconus) darkini Ro¨ckel, Korn & Richard, 1993 Conus (Embrikena) cebuensis Wils, 1990 Conus (Phasmoconus) daucus Hwass in Bruguie`re, 1792 Conus (Dauciconus) cedonulli Linnaeus, 1767 Conus (Stephanoconus) dayriti Ro¨ckel & da Motta, 1983 Conus (Splinoconus) centurio (Born, 1778) Conasprella (Kohniconus) decoratus Ro¨ckel, Rolań & Monteiro, 1980 Conus (Lautoconus) cepasi Trovaõ, 1975 Conus (Lautoconus) dedonderi (Goethaels & D. Monsecour, 2013) Conus (Strategoconus) ceruttii Cargile, 1997 Conus (Dauciconus) delanoyae Trovaõ, 1979 Conus (Lautoconus) cervus Lamarck, 1822 Conus (Textilia) delessertii (Rećluz, 1843) Conasprella (incertae chaldaeus (Ro¨ding, 1798) Conus (Virroconus) sedis) chiangi (Azuma, 1972) Conus (Stephanoconus) denizi (Afonso & Tenorio, 2011) Conus (Lautoconus) chiapponorum Lorenz, 2004 Conus (Textilia) derrubado Rolań & Fernandes, 1990 Conus (Lautoconus) chytreus Tryon, 1884 Conus (Lautoconus) desidiosus A. Adams, 1853 Conus (Lautoconus) ciderryi da Motta, 1985 Conus (Phasmoconus) deynzerorum Petuch, 1995 Conus (Dauciconus) cinereus Hwass in Bruguie`re, 1792 Conus (Phasmoconus) diadema G. B. Sowerby I, 1834 Conus (Lividoconus)

Continued Continued

16 CLASSIFICATION OF CONE SNAILS

Appendix Table 1. Continued Appendix Table 1. Continued

Species Genus (Subgenus) Species Genus (Subgenus) dianthus G. B. Sowerby III, 1882 Conus (Dauciconus) fijiensis (Moolenbeek, Ro¨ckel & Bouchet, 2008) Conasprella (Fusiconus) dictator (Melvill, 1898) Conasprella (Fusiconus) fijisulcatus Moolenbeek, Ro¨ckel & Bouchet, Conus (Asprella) diegoi (Cossignani, 2014) Conus (Lautoconus) 2008 dieteri (Moolenbeek, Zandbergen & Bouchet, Conasprella (Fusiconus) filmeri Rolań&Ro¨ckel, 2000 Conus (Lautoconus) 2008) fischoederi Ro¨ckel & da Motta, 1983 Conus (Pionoconus) diminutus Trovaõ & Rolań, 1986 Conus (Lautoconus) flamingo Petuch, 1980 Conus (Dauciconus) dispar G. B. Sowerby I, 1833 Conus (Dauciconus) flammeacolor Petuch, 1992 Conus (Dauciconus) distans Hwass in Bruguie`re, 1792 Conus (Fraterconus) flavescens G. B. Sowerby I, 1834 Conus (Dauciconus) dominicanus Hwass in Bruguie`re, 1792 Conus (Stephanoconus) flavidus Lamarck, 1810 Conus (Virgiconus) dondani (Kosuge, 1981) Profundiconus flavus Ro¨ckel, 1985 Conus (Phasmoconus) donnae Petuch, 1998 Conus (Dauciconus) flavusalbus Rolań&Ro¨ckel, 2000 Conus (Lautoconus) dorotheae Monnier & Limpalae¨r, 2010 Conus (Lautoconus) floccatus G. B. Sowerby I, 1841 Conus (Pionoconus) dorreensis Pe´ron, 1807 Conus (Virroconus) floridulus A. Adams & Reeve, 1848 Conus (Lividoconus) duffyi Petuch, 1992 Conus (Stephanoconus) fluviamaris (Petuch & Sargent, 2011) Conasprella (Ximeniconus) dusaveli (H. Adams, 1872) Conus (Textilia) fontonae Rolań&Trovaõ, 1990 Conus (Lautoconus)

ebraeus Linnaeus, 1758 Conus (Virroconus) fragilissimus Petuch, 1979 Conus (Gastridium) Downloaded from eburneus Hwass in Bruguie`re, 1792 Conus (Tesselliconus) franciscoi Rolań&Ro¨ckel, 2000 Conus (Lautoconus) echinophilus (Petuch, 1975) Conus (Lautoconus) frausseni (Tenorio & Poppe, 2004) Profundiconus echo Lauer, 1989 Conus (Darioconus) frigidus Reeve, 1848 Conus (Virgiconus) edaphus Dall, 1910 Conus (Tesselliconus) fulmen Reeve, 1843 Conus (Pionoconus) ednae (Petuch, 2013) Conus (Sandericonus) fumigatus Hwass in Bruguie`re, 1792 Conus (Splinoconus) edwardpauli Petuch, 1998 Conus (Dauciconus) furnae Rolań, 1990 Conus (Lautoconus) http://mollus.oxfordjournals.org/ eldredi Morrison, 1955 Conus (Gastridium) furvus Reeve, 1843 Conus (Calibanus) elegans (G. B. Sowerby III, 1895) Conasprella (Fusiconus) fuscoflavus Ro¨ckel, Rolań & Monteiro, 1980 Conus (Lautoconus) eleutheraensis (Petuch, 2013) Conus (Dauciconus) fuscolineatus G. B. Sowerby III, 1905 Conus (Lautoconus) elokismenos (Kilburn, 1975) Conasprella (Fusiconus) gabelishi da Motta & Ninomiya, 1982 Conus (Floraconus) emaciatus Reeve, 1849 Conus (Virgiconus) garciai da Motta, 1982 Conus (Dauciconus) emarginata (Reeve, 1844) Conasprella (Kohniconus) garywilsoni Lorenz & Morrison, 2004 Conus (Lividoconus) emersoni (G. D. Hanna, 1963) Profundiconus gauguini Richard & Salvat, 1973 Conus (Pionoconus) empressae Lorenz, 2001 Conus (Splinoconus) generalis Linnaeus, 1767 Conus (Strategoconus)

encaustus Kiener, 1845 Conus (Virroconus) genuanus Linnaeus, 1758 Conus (Kalloconus) by guest on December 3, 2014 episcopatus da Motta, 1982 Conus (Darioconus) geographus Linnaeus, 1758 Conus (Gastridium) ericmonnieri (Petuch & R. F. Myers, 2014) Conasprella (Ximeniconus) gibsonsmithorum Petuch, 1986 Conus (Dauciconus) ermineus Born, 1778 Conus (Chelyconus) gigasulcatus Moolenbeek, Ro¨ckel & Bouchet, Conus (Asprella) ernesti Petuch, 1990 Conus (Dauciconus) 2008 erythraeensis Reeve, 1843 Conus (Phasmoconus) gilberti (Bozzetti, 2012) Conus (Splinoconus) escondidai Poppe & Tagaro, 2005 Conus (Lividoconus) gilvus Reeve, 1849 Conus (Phasmoconus) estivali Moolenbeek & Richard, 1995 Conus (Klemaeconus) giorossii Bozzetti, 2005 Conus (Phasmoconus) eucoronata (G.B. Sowerby III, 1903) Conasprella (Conasprella) gladiator Broderip, 1833 Conus (Monteiroconus) eugrammata (Bartsch & Rehder, 1943) Conasprella (Conasprella) glans Hwass in Bruguie`re, 1792 Conus (Leporiconus) evansi Bondarev, 2001 Conus (Phasmoconus) glaucus Linnaeus, 1758 Conus (Dendroconus) eversoni Petuch, 1987 Conus (Attenuiconus) glenni Petuch, 1993 Conus (Atlanticonus) evorai Monteiro, Fernandes & Rolań, 1995 Conus (Lautoconus) glicksteini Petuch, 1987 Conus (Dauciconus) excelsus G. B. Sowerby III, 1908 Conus (Turriconus) gloriakiiensis Kuroda & Itoˆ, 1961 Conus (Splinoconus) exiguus Lamarck, 1810 Conus (Phasmoconus) gloriamaris Chemnitz, 1777 Conus (Cylinder) eximius Reeve, 1849 Conus (Lividoconus) glorioceanus Poppe & Tagaro, 2009 Conus (Cylinder) explorator Vink, 1990 Conus (Dauciconus) goajira Petuch, 1992 Conus (Dauciconus) fantasmalis Rolań, 1990 Conus (Lautoconus) gondwanensis Ro¨ckel & Moolenbeek, 1995 Conus (Turriconus) felitae Rolań, 1990 Conus (Lautoconus) gonsaloi (Afonso & Tenorio, 2014) Conus (Lautoconus) felix Fenzan, 2012 Conus (Plicaustraconus) gordyi (Ro¨ckel & Bondarev, 2000) Conasprella (Conasprella) fenzani (Petuch & Sargent, 2011) Conasprella (Kohniconus) goudeyi (Monnier & Limpalae¨r, 2012) Conus (Phasmoconus) fergusoni G. B. Sowerby II, 1873 Conus (Pyruconus) gradatulus Weinkauff, 1875 Conus (Sciteconus) fernandesi Tenorio, Afonso & Rolań, 2008 Conus (Lautoconus) gradatus W. Wood, 1828 Conus (Dauciconus) ferrugineus Hwass in Bruguie`re, 1792 Conus (Strategoconus) grahami Ro¨ckel, Cosel & Burnay, 1980 Conus (Lautoconus) fiadeiroi (Tenorio, Afonso, Cunha & Rolań, Conus (Lautoconus) granarius Kiener, 1847 Conus (Stephanoconus) 2014) grangeri G. B. Sowerby III, 1900 Conus (Asprella) figulinus Linnaeus, 1758 Conus (Dendroconus) granulatus Linnaeus, 1758 Conus (Atlanticonus)

Continued Continued

17 N. PUILLANDRE ET AL.

Appendix Table 1. Continued Appendix Table 1. Continued

Species Genus (Subgenus) Species Genus (Subgenus) granum Ro¨ckel & Fischo¨der, 1985 Conus (Leporiconus) jorioi (Petuch, 2013) Conus (Dauciconus) gratacapii Pilsbry, 1904 Conus (Turriconus) josephinae Rola´n, 1980 Conus (Lautoconus) grohi (Tenorio & Poppe, 2004) Conasprella joserochoi (Cossignani, 2014) Conus (Lautoconus) (Endemoconus) jourdani da Motta, 1984 Conus (Lautoconus) guanche Lauer, 1993 Conus (Lautoconus) jucundus G. B. Sowerby III, 1887 Conus (Dauciconus) gubernator Hwass in Bruguie`re, 1792 Conus (Pionoconus) judaeus Bergh, 1895 Conus (Virroconus) guidopoppei (G. Raybaudi Massilia, 2005) Conasprella (Fusiconus) julieandreae Cargile, 1995 Conus (Stephanoconus) guinaicus Hwass in Bruguie`re, 1792 Conus (Lautoconus) julii Lienard, 1870 Conus (Textilia) guyanensis (Van Mol, 1973) Conasprella (Dalliconus) kaiserae (Tenorio, Tucker & Chaney, 2012) Conus (Dauciconus) habui Lan, 2002 Conus (Phasmoconus) kalafuti da Motta, 1987 Conus (Dauciconus) hamamotoi Yoshiba & Koyama, 1984 Conus (Splinoconus) kanakinus (Richard, 1983) Profundiconus hamanni Fainzilber & Mienis, 1986 Conus (Leptoconus) kawamurai Habe, 1962 Conus (Leptoconus) hanshassi (Lorenz & Barbier, 2012) Conus (Strategoconus) kermadecensis Iredale, 1913 Conus (Lividoconus) harasewychi Petuch, 1987 Conus (Dauciconus) kersteni Tenorio, Afonso & Rola´n, 2008 Conus (Lautoconus) harlandi Petuch, 1987 Conus (Stephanoconus) kerstitchiWalls, 1978 Conus (Dauciconus)

havanensis Aguayo & Pe´rez Farfante, 1947 Conus (Dauciconus) kevani Petuch, 1987 Conus (Dauciconus) Downloaded from helgae Blo¨cher, 1992 Conus (Asprella) kiicumulus Azuma, 1982 Conus (Phasmoconus) henckesi (Coltro, 2004) Conasprella (Ximeniconus) kimioi (Habe, 1965) Conasprella hennequini Petuch, 1993 Conus (Dauciconus) (Boucheticonus) henriquei (Petuch & R. F. Myers, 2014) Conasprella (Coltroconus) kinoshitai Kuroda, 1956 Conus (Afonsoconus) herndli (Petuch & R. F. Myers, 2014) Conasprella (Ximeniconus) kintoki Habe & Kosuge, 1970 Conus (Virgiconus) hieroglyphus Duclos, 1833 Conus (Ductoconus) kirkandersi Petuch, 1987 Conus (Dauciconus) http://mollus.oxfordjournals.org/ hilli Petuch, 1990 Conus (Dauciconus) klemae (Cotton, 1953) Conus (Klemaeconus) hirasei (Kuroda, 1956) Conus (Klemaeconus) kohni (McLean & Nybakken, 1979) Conasprella (Kohniconus) hivanus Moolenbeek, Zandbergen & Bouchet, Conus (Strategoconus) korni G. Raybaudi Massilia, 1993 Conus (Pseudolilliconus) 2008 kostini Filmer, Monteiro, Lorenz & Verdasca, Conus (Embrikena) honkeri Petuch, 1988 Conus (Attenuiconus) 2012 honkerorum (Petuch & R. F. Myers, 2014) Conus (Dauciconus) koukae (Monnier, Limpalae¨r & Robin, 2013) Conus (Pionoconus) hopwoodi (Tomlin, 1937) Conasprella (Fusiconus) kremerorum Petuch, 1988 Conus (Dauciconus) howelli (Iredale, 1929) Conasprella kuiperi Moolenbeek, 2006 Conus (Pseudolilliconus)

(Endemoconus) kulkulcan Petuch, 1980 Conus (Dauciconus) by guest on December 3, 2014 hyaena Hwass in Bruguie`re, 1792 Conus (Rhizoconus) kuroharai (Habe, 1965) Conus (Phasmoconus) hybridus Kiener, 1847 Conus (Lautoconus) lamberti Souverbie, 1877 Conus (Darioconus) hypochlorus (Tomlin, 1937) Conasprella (Fusiconus) lani (Crandall, 1979) Profundiconus iansa (Petuch, 1979) Conasprella (Coltroconus) largilliertii Kiener, 1847 Conus (Dauciconus) ichinoseana (Kuroda, 1956) Conasprella (Fusiconus) laterculatus G. B. Sowerby II, 1870 Conus (Phasmoconus) ignotus Cargile, 1998 Conus (Kellyconus) laueri (Monnier & Limpalae¨r, 2013) Conus (Darioconus) ikedai (Ninomiya, 1987) Profundiconus lecourtorum (Lorenz, 2011) Conus (Virroconus) immelmani Korn, 1998 Conus (Leptoconus) leekremeri Petuch, 1987 Conus (Dauciconus) imperialis Linnaeus, 1758 Conus (Stephanoconus) legatus Lamarck, 1810 Conus (Cylinder) inconstans E. A. Smith, 1877 Conus (Dauciconus) lemniscatus Reeve, 1849 Conus (Lamniconus) inﬁnitus Rola´n, 1990 Conus (Lautoconus) lenavati da Motta & Ro¨ckel, 1982 Conus (Splinoconus) infrenatus Reeve, 1848 Conus (Sciteconus) lenhilli (Cargile, 1998) Conasprella (Dalliconus) inscriptus Reeve, 1843 Conus (Phasmoconus) lentiginosa (Reeve, 1844) Conasprella (Fusiconus) insculpta (Kiener, 1847) Conasprella (Fusiconus) leobottonii Lorenz, 2006 Conus (Pionoconus) iodostoma Reeve, 1843 Conus (Asprella) leobrerai da Motta & Martin, 1982 Conus (Phasmoconus) ione (Fulton, 1938) Conasprella leopardus (Ro¨ding, 1798) Conus (Lithoconus) (Endemoconus) levis (Bozzetti, 2012) Conus (Pseudolilliconus) irregularis G. B. Sowerby II, 1858 Conus (Lautoconus) levistimpsoni (Tucker, 2013) Conus (Dauciconus) isabelarum Tenorio & Afonso, 2004 Conus (Lautoconus) leviteni (Tucker, Tenorio & Chaney, 2011) Conus (Darioconus) jacarusoi Petuch, 1998 Conus (Dauciconus) lienardi Bernardi & Crosse, 1861 Conus (Phasmoconus) janowskyae (Tucker & Tenorio, 2011) Conasprella (Kohniconus) lightbourni Petuch, 1986 Conus (Bermudaconus) janus Hwass in Bruguie`re, 1792 Conus (Phasmoconus) limpusi Ro¨ckel & Korn, 1990 Conus (Eremiconus) jaspidea (Gmelin, 1791) Conasprella (Ximeniconus) lindae Petuch, 1987 Conus (Lindaconus) jeanmartini (G. Raybaudi Massilia, 1992) Profundiconus lineopunctatus Kaicher, 1977 Conus (Lautoconus) jickelii Weinkauff, 1873 Conus (Phasmoconus) lischkeanus Weinkauff, 1875 Conus (Lividoconus) joliveti (Moolenbeek, Ro¨ckel & Bouchet, 2008) Conasprella (Fusiconus) litoglyphus Hwass in Bruguie`re, 1792 Conus (Strategoconus)

Continued Continued

18 CLASSIFICATION OF CONE SNAILS

Appendix Table 1. Continued Appendix Table 1. Continued

Species Genus (Subgenus) Species Genus (Subgenus) litteratus Linnaeus, 1758 Conus (Elisaconus) moluccensis Ku¨ster, 1838 Conus (Phasmoconus) lividus Hwass in Bruguie`re, 1792 Conus (Lividoconus) monachus Linnaeus, 1758 Conus (Pionoconus) lizardensis Crosse, 1865 Conus (Eremiconus) moncuri Filmer, 2005 Conus (Elisaconus) lizarum (Raybaudi Massilia & da Motta, 1992) Conasprella (Fusiconus) monile Hwass in Bruguie`re, 1792 Conus (Strategoconus) lobitensis Kaicher, 1977 Conus (Lautoconus) monilifer Broderip, 1833 Conus (Dauciconus) locumtenens Blumenbach, 1791 Conus (Leptoconus) moolenbeeki Filmer, 2011 Conus (Phasmoconus) lohri Kilburn, 1972 Conus (Darioconus) mordeirae Rolań&Trovaõ, 1990 Conus (Lautoconus) longilineus Ro¨ckel, Rolań & Monteiro, 1980 Conus (Lautoconus) moreleti Crosse, 1858 Conus (Virgiconus) longurionis (Kiener, 1847) Conasprella (Fusiconus) morrisoni G. Raybaudi Massilia, 1991 Conus (Pionoconus) lorenzi Monnier & Limpalae¨r, 2012 Conasprella (Conasprella) mozambicus Hwass in Bruguie`re, 1792 Conus (Floraconus) loyaltiensis (Ro¨ckel & Moolenbeek, 1995) Profundiconus mucronatus Reeve, 1843 Conus (Phasmoconus) lozeti Richard, 1980 Conus (Plicaustraconus) mulderi Fulton, 1936 Conus (Phasmoconus) lucasi (Bozzetti, 2010) Conus (Textilia) muriculatus G. B. Sowerby I, 1833 Conus (Lividoconus) lucaya Petuch, 2000 Conus (Dauciconus) mus Hwass in Bruguie`re, 1792 Conus (Monteiroconus) luciae Moolenbeek, 1986 Conus (Turriconus) musicus Hwass in Bruguie`re, 1792 Conus (Harmoniconus)

lucidus (W. Wood, 1828) Conasprella (Ximeniconus) mustelinus Hwass in Bruguie`re, 1792 Conus (Rhizoconus) Downloaded from lugubris Reeve, 1849 Conus (Lautoconus) namocanus Hwass in Bruguie`re, 1792 Conus (Rhizoconus) luquei Rolań&Trovaõ, 1990 Conus (Lautoconus) nanus G. B. Sowerby I, 1833 Conus (Harmoniconus) luteus G. B. Sowerby I, 1833 Conus (Leporiconus) naranjus Trovaõ, 1975 Conus (Lautoconus) lynceus G. B. Sowerby II, 1858 Conus (Phasmoconus) natalaurantius (S. G. Veldsman, 2013) Conus (Darioconus) maculiferus G. B. Sowerby I, 1833 Conus (Dauciconus) natalis G. B. Sowerby II, 1858 Conus (Leptoconus) madagascariensis G. B. Sowerby II, 1858 Conus (Darioconus) navarroi Rolań, 1986 Conus (Lautoconus) http://mollus.oxfordjournals.org/ madecassinus (Bozzetti, 2012) Conus (Phasmoconus) negroides Kaicher, 1977 Conus (Lautoconus) magellanicus Hwass in Bruguie`re, 1792 Conus (Dauciconus) neptunus Reeve, 1843 Conus (Phasmoconus) magnificus Reeve, 1843 Conus (Darioconus) niederhoeferi (Monnier, Limpalae¨r & Lorenz, Conus (Phasmoconus) magnottei Petuch, 1987 Conus (Dauciconus) 2012) magus Linnaeus, 1758 Conus (Pionoconus) nielsenae Marsh, 1962 Conus (Splinoconus) mahogani (Reeve, 1843) Conasprella (Ximeniconus) nigromaculatus Ro¨ckel & Moolenbeek, 1992 Conus (Pionoconus) maioensis Trovaõ, Rolań&Fe´lix-Alves, 1990 Conus (Lautoconus) nigropunctatus G. B. Sowerby II, 1858 Conus (Pionoconus) malacanus Hwass in Bruguie`re, 1792 Conus (Splinoconus) nimbosus Hwass in Bruguie`re, 1792 Conus (Phasmoconus)

maldivus Hwass in Bruguie`re, 1792 Conus (Strategoconus) nobilis Linnaeus, 1758 Conus (Eugeniconus) by guest on December 3, 2014 mappa [Lightfoot], 1786 Conus (Stephanoconus) nobrei Trovaõ, 1975 Conus (Lautoconus) marchionatus Hinds, 1843 Conus (Eugeniconus) nocturnus [Lightfoot], 1786 Conus (Conus) marielae Rehder & Wilson, 1975 Conus (Phasmoconus) nodulosus G. B. Sowerby II, 1864 Conus (Cylinder) marmoreus Linnaeus, 1758 Conus (Conus) nucleus Reeve, 1848 Conus (Splinoconus) martensi E. A. Smith, 1884 Conus (Splinoconus) nussatella Linnaeus, 1758 Conus (Hermes) martinianus Reeve, 1844 Conus (Phasmoconus) nux Broderip, 1833 Conus (Harmoniconus) maya (Petuch & Sargent, 2011) Conus (Dauciconus) nybakkeni (Tenorio, Tucker & Chaney, 2012) Conus (Dauciconus) mazei (Deshayes, 1874) Conasprella (Dalliconus) obscurus G. B. Sowerby I, 1833 Conus (Gastridium) mcbridei Lorenz, 2005 Conus (Harmoniconus) ochroleucus Gmelin, 1791 Conus (Phasmoconus) mcgintyi (Pilsbry, 1955) Conasprella (Dalliconus) ogum (Petuch & R. F. Myers, 2014) Conasprella (Ximeniconus) medoci Lorenz, 2004 Conus (Dendroconus) oishii (Shikama, 1977) Conus (Phasmoconus) melissae Tenorio, Afonso & Rolań, 2008 Conus (Lautoconus) olgiatii Bozzetti, 2007 Conus (Splinoconus) melvilli G. B. Sowerby III, 1879 Conus (Quasiconus) omaria Hwass in Bruguie`re, 1792 Conus (Darioconus) memiae (Habe & Kosuge, 1970) Conasprella (Conasprella) orbignyi (Audouin, 1831) Conasprella (Fusiconus) mercator Linnaeus, 1758 Conus (Lautoconus) orion Broderip, 1833 Conus (Dauciconus) messiasi Rolań & Fernandes, 1990 Conus (Lautoconus) ortneri Petuch, 1998 Conus (Dauciconus) micropunctatus Rolań&Ro¨ckel, 2000 Conus (Lautoconus) ostrinus (Tucker & Tenorio, 2011) Conus (Dauciconus) miles Linnaeus, 1758 Conus (Rhizoconus) otohimeae (Kuroda & Itoˆ, 1961) Conasprella (Conasprella) milesi E. A. Smith, 1887 Conus (Turriconus) pacei (Petuch, 1987) Conasprella (Dalliconus) miliaris Hwass in Bruguie`re, 1792 Conus (Virroconus) pacificus (Moolenbeek & Ro¨ckel, 1996) Profundiconus milneedwardsi Jousseaume, 1894 Conus (Leptoconus) pagoda (Kiener, 1847) Conasprella (Conasprella) mindana (Hwass in Bruguie`re, 1792) Conasprella (Ximeniconus) papilliferus G. B. Sowerby I, 1834 Conus (Floraconus) miniexcelsus Olivera & Biggs, 2010 Conus (Turriconus) papuensis Coomans & Moolenbeek, 1982 Conus (Splinoconus) minnamurra (Garrard, 1961) Conus (Eremiconus) paraguana Petuch, 1987 Conus (Dauciconus) miruchae Ro¨ckel, Rolań & Monteiro, 1980 Conus (Lautoconus) parascalaris Petuch, 1987 Conus (Dauciconus) mitratus Hwass in Bruguie`re, 1792 Conus (Turriconus) parius Reeve, 1844 Conus (Phasmoconus)

Continued Continued

19 N. PUILLANDRE ET AL.

Appendix Table 1. Continued Appendix Table 1. Continued

Species Genus (Subgenus) Species Genus (Subgenus) parvatus Walls, 1979 Conus (Harmoniconus) rachelae (Petuch, 1988) Conasprella (Kohniconus) paschalli Petuch, 1998 Conus (Dauciconus) radiatus Gmelin, 1791 Conus (Phasmoconus) patae Abbott, 1971 Conus (Kellyconus) rainesae (McGinty, 1953) Conasprella (Dalliconus) patglicksteinae Petuch, 1987 Conus (Dauciconus) ranonganus da Motta, 1978 Conus (Phasmoconus) patriceae (Petuch & R. F. Myers, 2014) Conus (Lamniconus) raoulensis (Powell, 1958) Conasprella patricius Hinds, 1843 Conus (Pyruconus) (Endemoconus) paukstisi (Tucker, Tenorio & Chaney, 2011) Conus (Harmoniconus) rattus Hwass in Bruguie`re, 1792 Conus (Rhizoconus) paulae Petuch, 1988 Conus (Dauciconus) raulsilvai Rolań, Monteiro & Fernandes, 1998 Conus (Lautoconus) pauperculus G. B. Sowerby I, 1834 Conus (Splinoconus) rawaiensis da Motta, 1978 Conus (Rhizoconus) peli Moolenbeek, 1996 Conus (Rhizoconus) recluzianus Bernardi, 1853 Conus (Splinoconus) penchaszadehi Petuch, 1986 Conus (Dauciconus) recurvus Broderip, 1833 Conus (Dauciconus) pennaceus Born, 1778 Conus (Darioconus) reductaspiralis Walls, 1979 Conus (Splinoconus) pepeiu (Moolenbeek, Zandbergen & Bouchet, Conasprella (Fusiconus) regius Gmelin, 1791 Conus (Stephanoconus) 2008) regonae Rolań&Trovaõ, 1990 Conus (Lautoconus) pergrandis (Iredale, 1937) Conus (Embrikena) regularis G. B. Sowerby I, 1833 Conus (Dauciconus)

perplexa (G. B. Sowerby II, 1857) Conasprella (Ximeniconus) retifer Menke, 1829 Conus (Cylinder) Downloaded from pertusus Hwass in Bruguie`re, 1792 Conus (Rhizoconus) richardbinghami Petuch, 1993 Conus (Dauciconus) petergabrieli Lorenz, 2006 Conus (Phasmoconus) richardsae Ro¨ckel & Korn, 1992 Conus (Splinoconus) philippii Kiener, 1847 Conus (Dauciconus) richeri Richard & Moolenbeek, 1988 Conus (Phasmoconus) pica A. Adams & Reeve, 1848 Conus (Phasmoconus) ritae Petuch, 1995 Conus (Atlanticonus) pictus Reeve, 1843 Conus (Sciteconus) rizali Olivera & Biggs, 2010 Conus (Turriconus) pineaui Pin & Leung Tack, 1989 Conus (Lautoconus) roatanensis (Petuch & Sargent, 2011) Conasprella (Ximeniconus) http://mollus.oxfordjournals.org/ planorbis Born, 1778 Conus (Strategoconus) roberti (Richard, 2009) Conasprella (Dalliconus) plinthis Richard & Moolenbeek, 1988 Conus (Klemaeconus) robini (Limpalae¨r & Monnier, 2012) Conus (Pionoconus) polongimarumai Kosuge, 1980 Conus (Stephanoconus) roeckeli Rola´n, 1980 Conus (Lautoconus) poormani Berry, 1968 Conus (Dauciconus) rolani Ro¨ckel, 1986 Conus (Asprella) poremskii (Petuch & R. F. Myers, 2014) Conasprella (Ximeniconus) rosalindensis Petuch, 1998 Conus (Dauciconus) portobeloensis Petuch, 1990 Conus (Dauciconus) rosemaryae Petuch, 1990 Conus (Dauciconus) poulosi Petuch, 1993 Conus (Attenuiconus) roseorapum G. Raybaudi & da Motta, 1990 Conus (Splinoconus) praecellens A. Adams, 1855 Conus (Turriconus) rouxi (Monnier, Limpalae¨r & Robin, 2013) Conus (Pionoconus)

pretiosus G. Nevill & H. Nevill, 1874 Conus (Phasmoconus) royaikeni (S. G. Veldsman, 2010) Conus (Leptoconus) by guest on December 3, 2014 primus Ro¨ckel & Korn, 1990 Conus (Phasmoconus) rufimaculosus Macpherson, 1959 Conus (Klemaeconus) princeps Linnaeus, 1758 Conus (Ductoconus) ruthae (Monnier & Limpalae¨r, 2013) Conus (Splinoconus) priscai (Bozzetti, 2012) Conus (Cylinder) rutila (Menke, 1843) Conasprella (Parviconus) profundorum (Kuroda, 1956) Profundiconus saecularis (Melvill, 1898) Conasprella (Fusiconus) proximus G. B. Sowerby II, 1860 Conus (Phasmoconus) sagei (Korn & G. Raybaudi Massilia, 1993) Conasprella (Lilliconus) pseudaurantius Vink & Cosel, 1985 Conus (Stephanoconus) sahlbergi da Motta & Harland, 1986 Conus (Dauciconus) pseudimperialis Moolenbeek, Zandbergen & Conus (Stephanoconus) salletae (Cossignani, 2014) Conus (Lautoconus) Bouchet, 2008 salreiensis Rolań, 1980 Conus (Lautoconus) pseudocardinalis Coltro, 2004 Conus (Dauciconus) salzmanni G. Raybaudi Massilia & Rolań, 1997 Conus (Phasmoconus) pseudocuneolus Ro¨ckel, Rolań & Monteiro, Conus (Lautoconus) sanderi Wils & Moolenbeek, 1979 Conus (Sandericonus) 1980 sandwichensis Walls, 1978 Conus (Tesselliconus) pseudokimioi da Motta & Martin, 1982 Conasprella sanguineus Kiener, 1850 Conus (Stephanoconus) (Boucheticonus) sanguinolentus Quoy & Gaimard, 1834 Conus (Lividoconus) pseudonivifer Monteiro, Tenorio & Poppe, 2004 Conus (Kalloconus) santanaensis (Afonso & Tenorio, 2014) Conus (Lautoconus) pseudorbignyi (Ro¨ckel & Lan, 1981) Conasprella (Fusiconus) santinii (Monnier & Limpalae¨r, 2014) Conus (Phasmoconus) pulcher [Lightfoot], 1786 Conus (Kalloconus) saragasae Rolań, 1986 Conus (Lautoconus) pulcherrima Brazier, 1896 INCERTAE SEDIS sargenti (Petuch, 2013) Conasprella (Ximeniconus) (Kenyonia) sartii Korn, Niederho¨fer & Blo¨cher, 2002 Conus (Phasmoconus) pulicarius Hwass in Bruguie`re, 1792 Conus (Puncticulis) sauros (Garcia, 2006) Conasprella (Dalliconus) puncticulata (Hwass in Bruguie`re, 1792) Conasprella (Ximeniconus) sazanka Shikama, 1970 Conus (Splinoconus) purissimus Filmer, 2011 Conus (Splinoconus) scalaris Valenciennes, 1832 Conus (Dauciconus) purpurascens G. B. Sowerby I, 1833 Conus (Chelyconus) scalarispira (Bozzetti, 2012) Conus (Pseudolilliconus) pusio (Hwass in Bruguie`re, 1792) Conasprella (Ximeniconus) scalarissimus da Motta, 1988 Conus (Dauciconus) queenslandis da Motta, 1984 Conus (Splinoconus) scalptus Reeve, 1843 Conus (Phasmoconus) quercinus [Lightfoot], 1786 Conus (Lividoconus) scaripha (Dall, 1910) Conasprella (Kohniconus) quiquandoni Lorenz & Barbier, 2008 Conus (Phasmoconus) scopulicola (Okutani, 1972) Profundiconus

Continued Continued

20 CLASSIFICATION OF CONE SNAILS

Appendix Table 1. Continued Appendix Table 1. Continued

Species Genus (Subgenus) Species Genus (Subgenus) scopulorum Van Mol, Tursch & Kempf, 1971 Conus (Brasiliconus) tacomae Boyer & Pelorce, 2009 Conus (Lautoconus) scottjordani (Poppe, Monnier & Tagaro, 2012) Conus (Cylinder) taeniatus Hwass in Bruguie`re, 1792 Conus (Virroconus) sculletti Marsh, 1962 Conus (Papyriconus) tagaroae (Limpalae¨r & Monnier, 2013) Conus (Cylinder) sculpturatus Ro¨ckel & da Motta, 1986 Conus (Phasmoconus) taslei Kiener, 1850 Conus (Lautoconus) sennottorum Rehder & Abbott, 1951 Conus (Dauciconus) telatus Reeve, 1848 Conus (Cylinder) serranegrae Rolań, 1990 Conus (Lautoconus) tenuilineatus Rolań&Ro¨ckel, 2001 Conus (Lautoconus) sertacinctus Ro¨ckel, 1986 Conus (Phasmoconus) tenuistriatus G. B. Sowerby II, 1858 Conus (Leporiconus) shaskyi (Tenorio, Tucker & Chaney, 2012) Conus (Dauciconus) teodorae Rolań & Fernandes, 1990 Conus (Lautoconus) shikamai Coomans, Moolenbeek & Wils, 1985 Conus (Splinoconus) teramachii (Kuroda, 1956) Profundiconus sieboldii (Reeve, 1848) Conasprella terebra Born, 1778 Conus (Virgiconus) (Endemoconus) terryni Tenorio & Poppe, 2004 Conus (Pseudopterygia) silviae (Cossignani, 2014) Conus (Lautoconus) tessulatus Born, 1778 Conus (Tesselliconus) simonei (Petuch & R. F. Myers, 2014) Conasprella (Ximeniconus) tethys (Petuch & Sargent, 2011) Conus (Virgiconus) skoglundae (Tenorio, Tucker & Chaney, 2012) Conus (Dauciconus) textile Linnaeus, 1758 Conus (Cylinder) smirna (Bartsch & Rehder, 1943) Profundiconus thalassiarchus G. B. Sowerby I, 1834 Conus (Calibanus)

sogodensis (Poppe, Monnier & Tagaro, 2012) Conus (Phasmoconus) theodorei Petuch, 2000 Conus (Dauciconus) Downloaded from solangeae Bozzetti, 2004 Conus (Textilia) therriaulti (Petuch, 2013) Conus (Lindaconus) solidus Gmelin, 1791 Conus (Stephanoconus) thevenardensis da Motta, 1987 Conus (Splinoconus) solomonensis Delsaerdt, 1992 Conus (Phasmoconus) thomae Gmelin, 1791 Conus (Darioconus) somalica (Bozzetti, 2013) Conasprella tiaratus G. B. Sowerby I, 1833 Conus (Virroconus) (Endemoconus) tiki (Moolenbeek, Zandbergen & Bouchet, Conasprella (Fusiconus) spectrum Linnaeus, 1758 Conus (Phasmoconus) 2008) http://mollus.oxfordjournals.org/ sphacelatus G. B. Sowerby I, 1833 Conus (Dauciconus) timorensis Hwass in Bruguie`re, 1792 Conus (Textilia) spiceri Bartsch & Rehder, 1943 Conus (Virgiconus) tinianus Hwass in Bruguie`re, 1792 Conus (Floraconus) spiroﬁlis (Habe & Kosuge, 1970) Conasprella tirardi Ro¨ckel & Moolenbeek, 1996 Conus (Rhizoconus) (Endemoconus) tisii T. C. Lan, 1978 Conus (Embrikena) splendidulus G. B. Sowerby I, 1833 Conus (Strategoconus) tornata (G. B. Sowerby I, 1833) Conasprella (Ximeniconus) sponsalis Hwass in Bruguie`re, 1792 Conus (Harmoniconus) tostesi Petuch, 1986 Conus (Lamniconus) spurius Gmelin, 1791 Conus (Lindaconus) traceyi (Tucker & Stahlschmidt, 2010) Conasprella stahlschmidti (Tenorio & Tucker, 2014) Profundiconus (Pseudoconorbis)

stanfieldi Petuch, 1998 Conus (Dauciconus) traillii A. Adams, 1855 Conus (Pseudolilliconus) by guest on December 3, 2014 stearnsii (Conrad, 1869) Conasprella (Ximeniconus) traversiana (E. A. Smith, 1875) Conasprella (Fusiconus) stercusmuscarum Linnaeus, 1758 Conus (Pionoconus) trencarti Nolf & Verstraeten, 2008 Conus (Lautoconus) stimpsoni Dall, 1902 Conus (Dauciconus) tribblei Walls, 1977 Conus (Splinoconus) stocki (Coomans & Moolenbeek, 1990) Conasprella (Fusiconus) trigonus Reeve, 1848 Conus (Plicaustraconus) stramineus Lamarck, 1810 Conus (Phasmoconus) tristensis Petuch, 1987 Conus (Dauciconus) straturatus G. B. Sowerby II, 1865 Conus (Phasmoconus) trochulus Reeve, 1844 Conus (Kalloconus) striatellus Link, 1807 Conus (Strategoconus) troendlei Moolenbeek, Zandbergen & Bouchet, Conus (Splinoconus) striatus Linnaeus, 1758 Conus (Pionoconus) 2008 striolatus Kiener, 1848 Conus (Pionoconus) trovaoi Rolań&Ro¨ckel, 2000 Conus (Lautoconus) stupa (Kuroda, 1956) Conus (Turriconus) tuberculosus (Tomlin, 1937) Profundiconus stupella (Kuroda, 1956) Conus (Turriconus) tulipa Linnaeus, 1758 Conus (Gastridium) suduirauti Raybaudi Massilia, 2004 Conus (Strategoconus) tuticorinensis Ro¨ckel & Korn, 1990 Conus (Quasiconus) sugimotonis Kuroda, 1928 Conus (Klemaeconus) typhon Kilburn, 1975 Conus (Splinoconus) sukhadwalai Ro¨ckel & da Motta, 1983 Conus (Splinoconus) unifasciatus Kiener, 1850 Conus (Lautoconus) sulcatus Hwass in Bruguie`re, 1792 Conus (Asprella) urashimanus Kuroda & Itoˆ, 1961 Conus (Splinoconus) sulcocastaneus Kosuge, 1981 Conus (Asprella) vanhyningi (Rehder, 1944) Conasprella (Ximeniconus) sunderlandi Petuch, 1987 Conus (Dauciconus) vanvilstereni (Moolenbeek & Zandbergen, Conus (Splinoconus) suratensis Hwass in Bruguie`re, 1792 Conus (Dendroconus) 2013) sutanorcum Moolenbeek, Ro¨ckel & Bouchet, Conus (Phasmoconus) vappereaui Monteiro, 2009 Conus (Phasmoconus) 2008 variegatus Kiener, 1848 Conus (Lautoconus) suturatus Reeve, 1844 Conus (Tesselliconus) varius Linnaeus, 1758 Conus (Strategoconus) swainsoni Estival & Cosel, 1986 Conus (Strategoconus) vaubani (Ro¨ckel, Richard & Moolenbeek, 1995) Profundiconus swinneni (Tenorio, Afonso, Cunha & Rolań, Conus (Lautoconus) vautieri Kiener, 1847 Conus (Puncticulis) 2014) vayssierei Pallary, 1906 Conus (Lautoconus) sydneyensis G. B. Sowerby III, 1887 Conus (Mamiconus) velaensis Petuch, 1993 Conus (Dauciconus) tabidus Reeve, 1844 Conus (Monteiroconus) venezuelanus Petuch, 1987 Conus (Dauciconus)

Continued Continued

21 N. PUILLANDRE ET AL.

Appendix Table 1. Continued Appendix Table 2. Nominal genus-group names of Recent Conidae and their current status in the present classiﬁcation. Species Genus (Subgenus) Name Status ventricosus Gmelin, 1791 Conus (Lautoconus) venulatus Hwass in Bruguie`re, 1792 Conus (Kalloconus) Afonsoconus Tucker & Tenorio, 2013 Conus (Afonsoconus) verdensis Trova˜o, 1979 Conus (Lautoconus) Africonus Petuch, 1975 synonym of Conus (Lautoconus) vexillum Gmelin, 1791 Conus (Rhizoconus) ArubaconusPetuch, 2013 synonym of Conus (Ductoconus) victoriae Reeve, 1843 Conus (Cylinder) Asprella Schaufuss, 1869 Conus (Asprella) vicweei Old, 1973 Conus (Textilia) Atlanticonus Petuch & Sergent, 2012 Conus (Atlanticonus) vidua Reeve, 1843 Conus (Conus) Attenuiconus Petuch, 2013 Conus (Attenuiconus) vikingorum Petuch, 1993 Conus (Dauciconus) Austroconus Tucker & Tenorio, 2009 synonym of Conus (Mamiconus) villepinii P. Fischer & Bernardi, 1857 Conus (Dauciconus) Bathyconus Tucker & Tenorio, 2009 synonym of Conasprella viminea (Reeve, 1849) Conasprella (Fusiconus) (Fusiconus) viola Cernohorsky, 1977 Conus (Splinoconus) Bermudaconus Petuch, 2013 Conus (Bermudaconus) violaceus Gmelin, 1791 Conus (Virgiconus) Boucheticonus Tucker & Tenorio, 2013 Conasprella (Boucheticonus) virgatus Reeve, 1849 Conus (Dauciconus) Brasiliconus Petuch, 2013 Conus (Brasiliconus) virgo Linnaeus, 1758 Conus (Virgiconus) Calamiconus Tucker & Tenorio, 2009 synonym of Conus (Lividiconus) visagenus Kilburn, 1974 Conus (Plicaustraconus) Calibanus da Motta, 1991 Conus (Calibanus) Californiconus Tucker & Tenorio, 2009 Californiconus visseri Delsaerdt, 1990 Conus (Pseudolilliconus) Downloaded from vittatus Hwass in Bruguie`re, 1792 Conus (Dauciconus) Cariboconus Petuch, 2003 synonym of Conus (Dauciconus) voluminalis Reeve, 1843 Conus (Splinoconus) Chelyconus Mo¨rch, 1852 Conus (Chelyconus) vulcanus Tenorio & Afonso, 2004 Conus (Lautoconus) Cleobula Iredale, 1930 synonym of Conus (Dendroconus) wakayamaensis (Kuroda, 1956) Conasprella (Conasprella) Coltroconus Petuch, 2013 Conasprella (Coltroconus) wallacei (Lorenz & Morrison, 2004) Conus (Pseudolilliconus) Conasprella Thiele, 1929 Conasprella (Conasprella) wallangra (Garrard, 1961) Conus (Plicaustraconus) Conasprelloides Tucker & Tenorio, 2009 synonym of Conus (Dauciconus) http://mollus.oxfordjournals.org/ wandae (Cossignani, 2014) Conus (Lautoconus) Continuconus Tucker & Tenorio, 2013 synonym of Conus (Klemaeconus) wendrosi (Tenorio & Afonso, 2013) Conasprella (Ximeniconus) Conus Linnaeus, 1758 Conus (Conus) wilsi Delsaerdt, 1998 Conus (Rhizoconus) Cornutoconus Suzuki, 1972 synonym of Conus (Taranteconus) wittigi Walls, 1977 Conus (Phasmoconus) Coronaxis Swainson, 1840 synonym of Conus (Conus) xanthicus Dall, 1910 Conus (Dauciconus) Cucullus Ro¨ding, 1798 synonym of Conus (Conus) xanthocinctus Petuch, 1986 Conus (Lamniconus) Cylinder Montfort, 1810 synonym of Conus (Cylindrus) xicoi Ro¨ckel, 1987 Conus (Lautoconus) Cylindrella Swainson, 1840 synonym of Conus (Asprella) ximenes (Gray, 1839) Conasprella (Ximeniconus) Cylindrus Batsch, 1789 Conus (Cylindrus) Dalliconus Tucker & Tenorio, 2009 Conasprella (Dalliconus)

yemenensis Bondarev, 1997 Conus (Phasmoconus) by guest on December 3, 2014 zandbergeni Filmer & Moolenbeek, 2010 Conus (Phasmoconus) Darioconus Iredale, 1930 Conus (Darioconus) zapatosensis Ro¨ckel, 1987 Conus (Phasmoconus) Dauciconus Cotton, 1945 Conus (Dauciconus) zebra Lamarck, 1810 Conus (Phasmoconus) Dendroconus Swainson, 1840 Conus (Dendroconus) zebroides Kiener, 1848 Conus (Lautoconus) Ductoconus da Motta, 1991 Conus (Ductoconus) zeylanicus Gmelin, 1791 Conus (Puncticulis) Duodenticonus Tucker & Tenorio, 2013 synonym of Conasprella ziczac Mu¨hlfeld, 1816 Conus (Dauciconus) (Conasprella) zinhoi (Cossignani, 2014) Conus (Lautoconus) Dyraspis Iredale, 1949 synonym of Conus (Virroconus) zonatus Hwass in Bruguie`re, 1792 Conus (Stephanoconus) Elisaconus Tucker & Tenorio, 2013 Conus (Elisaconus) zylmanae Petuch, 1998 Conus (Dauciconus) Embrikena Iredale, 1937 Conus (Embrikena) Endemoconus Iredale, 1931 Conasprella (Endemoconus) Eremiconus Tucker & Tenorio, 2009 Conus (Eremiconus) Erythroconus da Motta, 1991 synonym of Conus (Darioconus) Eugeniconus da Motta, 1991 Conus (Eugeniconus) Floraconus Iredale, 1930 Conus (Floraconus) Fraterconus Tucker & Tenorio, 2013 Conus (Fraterconus) Fulgiconus da Motta, 1991 synonym of Conus (Phasmoconus) Fumiconus da Motta, 1991 synonym of Conasprella (Fusiconus) Fusiconus da Motta, 1991 Conasprella (Fusiconus) Gastridium Modeer, 1793 Conus (Gastridium) Genuanoconus Tucker & Tenorio, 2009 synonym of Conus (Kalloconus) Gladioconus Tucker & Tenorio, 2009 synonym of Conus (Monteiroconus) Globiconus Tucker & Tenorio, 2009 synonym of Conasprella (Ximeniconus) Gradiconus da Motta, 1991 synonym of Conus (Dauciconus) Graphiconus da Motta, 1991 synonym of Conus (Phasmoconus)

Continued

22 CLASSIFICATION OF CONE SNAILS

Appendix Table 2. Continued Appendix Table 2. Continued

Name Status Name Status

Harmoniconus da Motta, 1991 Conus (Harmoniconus) Quasiconus Tucker & Tenorio, 2009 Conus (Quasiconus) Hermes Montfort, 1810 Conus (Hermes) Regiconus Iredale, 1930 synonym of Conus (Darioconus) Heroconus da Motta, 1991 synonym of Conus (Pionoconus) Rhizoconus Mo¨rch, 1852 Conus (Rhizoconus) Isoconus Tucker & Tenorio, 2013 synonym of Conus (Splinoconus) Rhombiconus Tucker & Tenorio, 2009 synonym of Conus Jaspidiconus Petuch, 2003 synonym of Conasprella (Stephanoconus) (Ximeniconus) Rhombus Montfort, 1810 synonym of Conus (Rhombiconus) Kalloconus da Motta, 1991 Conus (Kalloconus) Rolaniconus Tucker & Tenorio, 2009 synonym of Conus (Strategoconus) Kellyconus Petuch, 2013 Conus (Kellyconus) Rollus Montfort, 1810 synonym of Conus (Gastridium) Kenyonia Brazier, 1896 Conus (Kenyonia) Rubroconus Tucker & Tenorio, 2013 Conus (Rubroconus) Kermasprella Powell, 1958 synonym of Conasprella Sandericonus Petuch, 2013 Conus (Sandericonus) (Endemoconus) Sciteconus da Motta, 1991 Conus (Sciteconus) Ketyconus da Motta, 1991 synonym of Conus (Floraconus) Seminoleconus Petuch, 2003 synonym of Conus (Protoconus) Kioconus da Motta, 1991 synonym of Conus (Splinoconus) Socioconus da Motta, 1991 synonym of Conus (Pionoconus) Klemaeconus Tucker & Tenorio, 2013 Conus (Klemaeconus) Splinoconus da Motta, 1991 Conus (Splinoconus) Kohniconus Tucker & Tenorio, 2009 Conasprella (Kohniconus) Spuriconus Petuch, 2003 synonym of Conus (Lindaconus)

Kurodaconus Shikama & Habe, 1968 synonym of Conus (Turriconus) Stellaconus Tucker & Tenorio, 2009 synonym of Conus (Splinoconus) Downloaded from Lamniconus da Motta, 1991 Conus (Lamniconus) Stephanoconus Mo¨rch, 1852 Conus (Stephanoconus) Lautoconus Monterosato, 1923 Conus (Lautoconus) Strategoconus da Motta, 1991 Conus (Strategoconus) Leporiconus Iredale, 1930 Conus (Leporiconus) Strioconus Thiele, 1929 synonym of Conus (Pionoconus) Leptoconus Swainson, 1840 Conus (Leptoconus) Sulciconus Bielz, 1869 synonym of Conus (Asprella) Lilliconus Raybaudi Massilia, 1994 Conasprella (Lilliconus) Taranteconus Azuma, 1972 synonym of Conus Lindaconus Petuch, 2002 Conus (Lindaconus) (Stephanoconus) http://mollus.oxfordjournals.org/ Lithoconus Mo¨rch, 1852 Conus (Lithoconus) Tenorioconus Petuch & Drohlshagen, synonym of Conus Lividoconus Wils, 1970 Conus (Lividoconus) 2011 (Stephanoconus) Lizaconus da Motta, 1991 synonym of Conus (Profundiconus) Tesselliconus da Motta, 1991 Conus (Tesselliconus) Magelliconus da Motta, 1991 synonym of Conus (Purpuriconus) Textilia Swainson, 1840 Conus (Textilia) Mamiconus Cotton & Godfrey, 1932 Conus (Mamiconus) Thalassiconus Tucker & Tenorio, 2013 synonym of Conus (Calibanus) Miliariconus Tucker & Tenorio, 2009 synonym of Conus (Virroconus) Theliconus Swainson, 1840 synonym of Conus (Hermes) Mitraconus Tucker & Tenorio, 2013 synonym of Conus (Turriconus) Thoraconus da Motta, 1991 synonym of Conus (Fulgiconus) Monteiroconus da Motta, 1991 Conus (Monteiroconus) Trovaoconus Tucker & Tenorio, 2009 synonym of Conus (Kalloconus)

Nataliconus Tucker & Tenorio, 2009 synonym of Conus (Leptoconus) Tuckericonus Petuch, 2013 synonym of Conus (Dauciconus) by guest on December 3, 2014 Nimboconus Tucker & Tenorio, 2013 synonym of Conus (Phasmoconus) Tuliparia Swainson, 1840 synonym of Conus (Gastridium) Nitidoconus Tucker & Tenorio, 2013 synonym of Conus (Splinoconus) Turriconus Shikama & Habe, 1968 Conus (Turriconus) Ongoconus da Motta, 1991 synonym of Conus (Kioconus) Utriculus Schumacher, 1817 synonym of Conus (Gastridium) Papyriconus Tucker & Tenorio, 2013 Conus (Papyriconus) Varioconus da Motta, 1991 synonym of Conus (Lautoconus) Parviconus Cotton & Godfrey, 1932 Conasprella (Parviconus) Viminiconus Tucker & Tenorio, 2009 synonym of Conasprella Perplexiconus Tucker & Tenorio, 2009 synonym of Conasprella (Fusiconus) (Ximeniconus) Virgiconus Cotton, 1945 Conus (Virgiconus) Phasmoconus Mo¨rch, 1852 Conus (Phasmoconus) Virroconus Iredale, 1930 Conus (Virroconus) Pionoconus Mo¨rch, 1852 Conus (Pionoconus) Vituliconus da Motta, 1991 synonym of Conus (Strategoconus) Plicaustraconus Moolenbeek, 2008 Conus (Plicaustraconus) Ximeniconus Emerson & Old, 1962 Conasprella (Ximeniconus) Poremskiconus Petuch, 2013 synonym of Conus (Dauciconus) Yeddoconus Tucker & Tenorio, 2009 synonym of Conasprella Profundiconus Kuroda, 1956 Profundiconus (Endemoconus) Protoconus da Motta, 1991 synonym of Conus (Tenorioconus) Protostrioconus Tucker & Tenorio, 2009 synonym of Conus (Gastridium) Pseudoconorbis Tucker & Tenorio, Conasprella (Pseudoconorbis) 2009 Pseudohermes Tucker & Tenorio, 2013 synonym of Conus (Virgiconus) Pseudolilliconus Tucker & Tenorio, Conus (Pseudolilliconus) 2009 Pseudonoduloconus Tucker & Tenorio, Conus (Pseudonoduloconus) 2009 Pseudopterygia Tucker & Tenorio, Conus (Pseudopterygia) 2013 Puncticulis Swainson, 1840 Conus (Puncticulis) Purpuriconus da Motta, 1991 synonym of Conus (Dauciconus) Pyruconus Olsson, 1967 Conus (Pyruconus)

Continued