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6. Selected Paleozoic (Carboniferous/Permian) caenogastropods and putative outgroup taxa. (A, B) Protoconch of a naticopsid (Neritimorpha) from the Mississippian (Lower Carboniferous, ca. 330 Mya; Ruddle Shale, Arkansas, United States). Protoconchs of Recent neritimorphs are highly convolute with resorbed inner whorls; however, protoconchs of Naticopsidae show no major differences from those of caenogastropods, except for a relatively high whorl expansion rate. Width 0.8 mm (from Nützel and Mapes 2001). (C, D) Cerithimorph caenogastropod from the Upper Carboniferous (Late Moscovian, c. 305 Mya; Buckhorn Asphalt deposit, Oklahoma, United States); this small heliciform, planktotrophic larval shell resembles the protoconch of some modern cerithioids. C, height 2.0 mm; D, height 0.4 mm. (E–G) Stegocoelia (Goniasmatidae, Palaeostyloidea), a slit-bearing caenogastropod from the Upper Carboniferous (Late Moscovian, c. 305 Mya; Buckhorn Asphalt deposit, Oklahoma, United States), representative of a rich late Paleozoic group of Murchisonia resembling caenogastropods; E, teleoconch detail showing slitlike structure (selenizone) slightly above mid-whorl; width 0.5 mm; F, height 2.0 mm; G, protoconch in side view, a lecithotrophic larval shell with a distinct sinusigera. Protoconchs of planktotrophic species of this group resemble Figure 13.6D; height 0.36 mm (from Bandel et al . 2002). (H) Soleniscus , a widespread subulitoid (Soleniscidae) from the Upper Carboniferous (Gzhelian, c. 300 Mya; Finis Shale, Texas, United States), showing a distinct, twisted siphonal canal, a columellar fold, and a smooth larval shell; height 3.2 mm (from Nützel et al . 2000). (I–K) Imogloba (Imoglobidae) from the Mississippian (Lower Carboniferous, c. 330 Mya of Arkansas, United States); these globular, subulitoid gastropods have an open coiled initial whorl followed by early whorls (probably larval shell) with a very characteristic ornament of noncollabral threads; I, height 2.5 mm; J, probably isolated larval shell, height 0.85 mm; K, width 4.4 mm. (L, M) Pseudozygopleura (Pseudozygopleuridae) from the Late Carboniferous (ca. 300 Mya, Gzhelian, Ames Shale, West Virginia, United States); pseudozygopleurids were abundant and diverse for about 100 million years (during the late Paleozoic) and became extinct at the end-Permian mass extinction event; they have highly characteristic larval shells with an ornament of curving, collabral ribs that form a spiral thread (from Nützel 1998); L, larval shell; height 0.86 mm; M, height 3.0 mm. 

6. Selected Paleozoic (Carboniferous/Permian) caenogastropods and putative outgroup taxa. (A, B) Protoconch of a naticopsid (Neritimorpha) from the Mississippian (Lower Carboniferous, ca. 330 Mya; Ruddle Shale, Arkansas, United States). Protoconchs of Recent neritimorphs are highly convolute with resorbed inner whorls; however, protoconchs of Naticopsidae show no major differences from those of caenogastropods, except for a relatively high whorl expansion rate. Width 0.8 mm (from Nützel and Mapes 2001). (C, D) Cerithimorph caenogastropod from the Upper Carboniferous (Late Moscovian, c. 305 Mya; Buckhorn Asphalt deposit, Oklahoma, United States); this small heliciform, planktotrophic larval shell resembles the protoconch of some modern cerithioids. C, height 2.0 mm; D, height 0.4 mm. (E–G) Stegocoelia (Goniasmatidae, Palaeostyloidea), a slit-bearing caenogastropod from the Upper Carboniferous (Late Moscovian, c. 305 Mya; Buckhorn Asphalt deposit, Oklahoma, United States), representative of a rich late Paleozoic group of Murchisonia resembling caenogastropods; E, teleoconch detail showing slitlike structure (selenizone) slightly above mid-whorl; width 0.5 mm; F, height 2.0 mm; G, protoconch in side view, a lecithotrophic larval shell with a distinct sinusigera. Protoconchs of planktotrophic species of this group resemble Figure 13.6D; height 0.36 mm (from Bandel et al . 2002). (H) Soleniscus , a widespread subulitoid (Soleniscidae) from the Upper Carboniferous (Gzhelian, c. 300 Mya; Finis Shale, Texas, United States), showing a distinct, twisted siphonal canal, a columellar fold, and a smooth larval shell; height 3.2 mm (from Nützel et al . 2000). (I–K) Imogloba (Imoglobidae) from the Mississippian (Lower Carboniferous, c. 330 Mya of Arkansas, United States); these globular, subulitoid gastropods have an open coiled initial whorl followed by early whorls (probably larval shell) with a very characteristic ornament of noncollabral threads; I, height 2.5 mm; J, probably isolated larval shell, height 0.85 mm; K, width 4.4 mm. (L, M) Pseudozygopleura (Pseudozygopleuridae) from the Late Carboniferous (ca. 300 Mya, Gzhelian, Ames Shale, West Virginia, United States); pseudozygopleurids were abundant and diverse for about 100 million years (during the late Paleozoic) and became extinct at the end-Permian mass extinction event; they have highly characteristic larval shells with an ornament of curving, collabral ribs that form a spiral thread (from Nützel 1998); L, larval shell; height 0.86 mm; M, height 3.0 mm. 

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Caenogastropods comprise about 60% of living gastropod species and include a large number of ecologically and commercially important marine families. They have undergone an extraordinary adaptive radiation, resulting in considerable morphological, ecological, physiological , and behavioral diversity. There is a wide array of often convergent shell...

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... As noted by Ponder et al. (2008), triphoroideans have remained virtually untouched by comparative approaches. The anatomy of Pseudomohnia agrees well with what little has been described for members of the superfamily thus far (Risbec 1943;Fretter 1951;Kosuge 1966;Houston 1985), and for newtoniellids in particular (e.g., Houbrick 1987;Golding et al. 2009), and in their ecology as feeders on sponges. ...
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Mohnia kurilana Dall, 1913 was described more than 100 years ago from deep waters off the Kuril Islands and remains exceedingly rare in museum collections. Originally placed in the carnivorous neogastropod family Buccinidae, fragmentary soft parts from the type lot and from several specimens belonging to allied species collected in the Aleutian Islands in the 1990s have allowed anatomical investigations for the first time. These have revealed the presence of a paucispiral operculum with an eccentric nucleus, foot with a deep propodial pedal gland and metapodial pedal gland, taenioglossate radula, short acrembolic proboscis, well-developed mid-esophageal gland, glandular prostate, and the absence of a penis; the nervous system is epiathroid with a long supra-esophageal connective and numerous statoconia in the statocysts. Analysis of the gut contents revealed abundant halichondriid sponge spicules. This evidence indicates a placement in the Triphoroidea, a diverse superfamily of specialized spongivores. Mohnia kurilana is transferred to the Newtoniellidae and placed in the new genus Pseudomohnia gen. nov. Pseudomohnia rogerclarki sp. nov. is established for a new species from the Aleutian Islands characterized by its narrowly turreted shell and distinctive multicuspid rachidian. A lectotype is designated for Mohnia kurilana .
... Equally controversial is the higher phylogenetic placement of the superfamily. Traditionally perceived as "the most advanced superfamily of the mesogastropods" (Houbrick & Fretter, 1969: 415), morphological approaches to caenogastropod phylogeny inevitably place the Tonnoidea as sister to the Neogastropoda (e.g., Ponder et al., 2008;Simone, 2011), while molecular approaches typically resolve them among the neogastropods (e.g., Riedel, 2000;Colgan et al., 2007;Ponder et al., 2008;Osca et al., 2015). However, representation of Tonnoidea in gastropod phylogenies has been very sparse and has not allowed the circumscription of the superfamily and of its constituent families to be assessed robustly. ...
... Equally controversial is the higher phylogenetic placement of the superfamily. Traditionally perceived as "the most advanced superfamily of the mesogastropods" (Houbrick & Fretter, 1969: 415), morphological approaches to caenogastropod phylogeny inevitably place the Tonnoidea as sister to the Neogastropoda (e.g., Ponder et al., 2008;Simone, 2011), while molecular approaches typically resolve them among the neogastropods (e.g., Riedel, 2000;Colgan et al., 2007;Ponder et al., 2008;Osca et al., 2015). However, representation of Tonnoidea in gastropod phylogenies has been very sparse and has not allowed the circumscription of the superfamily and of its constituent families to be assessed robustly. ...
... Higher order phylogenetic analyses of caenogastropods typically have included only one or two representative tonnoideans (e.g., Colgan et al., 2007;Ponder et al., 2008;Osca et al., 2015); no molecular investigation has included members of both Ficoidea and Tonnoidea except that of Riedel (2000). The only analysis with sufficient sampling to assess monophyly and relationships among tonnoidean families is that of Simone (2011) based on morphological data. ...
Article
The Tonnoidea is a moderately diverse group of large, predatory gastropods with ∼360 valid species. Known for their ability to secrete sulfuric acid, they use it to prey on a diversity of invertebrates, primarily echinoderms. Tonnoideans currently are classified in seven accepted families: the comparatively well known, shallow water Bursidae, Cassidae, Personidae, Ranellidae, and Tonnidae, and the lesser-known, deep water Laubierinidae and Pisanianuridae. We assembled a mitochondrial and nuclear gene (COI, 16S, 12S, 28S) dataset for ∼80 species and 38 genera currently recognized as valid. Bayesian analysis of the concatenated dataset recovered a monophyletic Tonnoidea, with Ficus as its sister group. Unexpectedly, Thalassocyon, currently classified in the Ficidae, was nested within the ingroup as the sister group to Distorsionella. Among currently recognized families, Tonnidae, Cassidae, Bursidae and Personidae were supported as monophyletic but the Ranellidae and Ranellinae were not, with Cymatiinae, Ranella and Charonia supported as three unrelated clades. The Laubierinidae and Pisanianuridae together form a monophyletic group. Although not all currently accepted genera have been included in the analysis, the new phylogeny is sufficiently robust and stable to the inclusion/exclusion of nonconserved regions to establish a revised family-level classification with nine families: Bursidae, Cassidae, Charoniidae, Cymatiidae, Laubierinidae, Personidae, Ranellidae, Thalassocyonidae and Tonnidae. The results reveal that many genera as presently circumscribed are para- or polyphyletic and, in some cases support the rescue of several genus-group names from synonymy (Austrosassia, Austrotriton, Laminilabrum, Lampadopsis, Personella, Proxicharonia, Tritonoranella) or conversely, support their synonymization (Biplex with Gyrineum). Several species complexes are also revealed that merit further investigation (e.g., Personidae: Distorsio decipiens, D. reticularis; Bursidae: Bursa tuberosissima; Cassidae: Echinophoria wyvillei, Galeodea bituminata, and Semicassis bisulcata). Consequently, despite their teleplanic larvae, the apparently circumglobal distribution of some tonnoidean species is the result of excessive synonymy. The superfamily is estimated to have diverged during the early Jurassic (∼186 Ma), with most families originating during a narrow ∼20 My window in Albian-Aptian times as part of the Mesozoic Marine Revolution.
... Digestive, circulatory, excretory, reproductive and central nervous systems were investigated in detail. The terminology used for odontophore muscles was based on Ponder et al. (2008), Simone (2011) and Lima and Simone (2015). Drawings were done with the aid of a camera lucida. ...
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The last review of the genus Actinocyclus consider only two valid species for the genus: Actinocyclusverrucosus Ehrenberg, 1831 (type species of the genus) and Actinocycluspapillatus (Bergh, 1878), both with a geographical distribution in the Indo-Pacific. The anatomy of these species is still unknown, except for some scanty anatomical information. A detailed anatomical study of Actinocyclusverrucosus is performed, including inedited structures such as digestive system, odontophore muscles and circulatory system, beyond complementary information on the commonly studied structures, in order to clarify the taxonomy and distribution.
... Since there is doubt whether the Caribbean and Brazilian populations of Platydoris angustipes are the same nervous systems were investigated in detail. The terminology used for odontophore muscles was based on Ponder et al. (2008), Simone (2011) and Lima and Simone (2015). Digital photographs were taken at each dissection step. ...
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Platydoris angustipes (Mörch, 1863) is a common nudibranch in the Western Atlantic, ranging from Florida, USA, to Rio de Janeiro, Brazil. In this study, we examined the anatomy of P. angustipes along its distribution, including its type material. Our analysis shows consistent differences between the Caribbean and Brazilian populations, mainly in the reproductive system, radular teeth and odontophore musculature. This strongly suggests that the two populations actually belong to distinct species. The Brazilian population is described herein as a new species, Platydoris guarani sp. n.
... For transmission electron microscopy (TEM), the gonads of eight individuals were examined. After removal from the shell, small Table 1 Morphology of the sperm and parasperm papers published on neogastropod species (Ponder et al., 2008 pieces of the testis were fixed in 2.5% glutaraldehyde in 0.1 M sodium phosphate buffer containing filtered seawater for 4 h at 48C, and washed thoroughly either in sucrose-adjusted cacodylate buffer or in CaCl 2 -adjusted phosphate buffer. Subsequently, the tissue pieces were placed in a 1% solution of osmium tetroxide (in 0.1 M cacodylate or phosphate buffer) for 1.5 h, and again washed in buffer. ...
... The superfamily Triphoroidea Gray comprises the marine families Triphoridae Gray, Cerithiopsidae Adams and Adams and Newtoniellidae Korobkov (Ponder and Bouchet 2005), although relationships within the superfamily and its phylogenetic position within Caenogastropoda remain unresolved (Ponder et al. 2008). The distinctive morphology of triphoroideans, exemplified by their high-spired shells, might be explained by their ancient isolation from other gastropod lineages (Nützel 1998;Ponder et al. 2008). ...
... The superfamily Triphoroidea Gray comprises the marine families Triphoridae Gray, Cerithiopsidae Adams and Adams and Newtoniellidae Korobkov (Ponder and Bouchet 2005), although relationships within the superfamily and its phylogenetic position within Caenogastropoda remain unresolved (Ponder et al. 2008). The distinctive morphology of triphoroideans, exemplified by their high-spired shells, might be explained by their ancient isolation from other gastropod lineages (Nützel 1998;Ponder et al. 2008). They are primarily sponge feeders (Wells 1998), and their adaptation to a micropredatory lifestyle has led to a rapid evolution of the radula and an enormous morphological diversity (Nützel 1998), correlated to the high level of variation in structural and textural properties of sponges (Marshall 1978). ...
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The marine gastropod superfamily Triphoroidea is composed of three families: Triphoridae and Cerithiopsidae are usually associated with shallow waters, and Newtoniellidae is historically associated with the deep sea. We provide an extensive review of the literature and delineate the groups within the three families that are truly linked to the deep sea, with reference to their developmental mode. Two genera of Triphoridae, Strobiligera and Inella, are commonly found in deep waters, and Metaxia presents some species with apparently wide depth ranges. Some deep-sea genera currently placed in Cerithiopsidae, such as Ektonos and Krachia, are very similar to Newtoniellidae; a phylogeny of the superfamily is needed to clarify the distinction of the two families. The majority of Triphoroidea species from the deep sea undergo non-planktotrophic development, although planktotrophy is not uncommon.
... 2) with Hebeulima (Vanikoroidea: Eulimidae) as a sister to the rissoinid-barleeiid clade. A rissoinid-eulimid clade was retrieved in nearly all previous caenogastropod phylogenies (Colgan et al., 2007;Ponder et al., 2008). However, those studies included no other rissooidean family. ...
Article
The Rissooidea is an evolutionarily ancient and mega-diverse group of marine micro-gastropods that occur from intertidal to deep waters at all latitudes. Their current systematics is predominantly based on phenetic grounds and there has been no comprehensive molecular phylogeny. Based on sequences of mitochondrial and nuclear DNA from the most complete sampling of Rissoidae to date, this work represents the first treatment of the group performed through a phylogenetic approach. The main goals are to clarify the phylogenetic position of the Rissoidae, investigate the relationships within rissoid taxa and test the utility of some diagnostic morphological traits. Our phylogeny indicates that the Rissoidae are one of six distinct family-lineages within the superfamily Rissooidea (along with Barleeiidae, Emblandidae, Lironobidae, Rissoinidae and Zebinidae) whose recognition is supported by several synapomorphies. While most of the characters studied exhibit widespread convergence, some others prove useful in separating genera and broader taxonomic groups. The relationships among rissoid taxa challenge the current systematics, indicating the non-monophyly of some genera with purportedly transoceanic distribution and the need of taxonomic revision for some highly diverse genera. Our phylogeny suggests that the Rissoidae originated in shallow seas and independently radiated into bathyal waters at least twice.
... With thousands of described species, the caenogastropods (periwinkles, whelks, cones, cowries, miters, etc.) constitute the largest and most evolutionary successful group of living gastropods (Ponder et al., 2008). These snails are highly diverse in the morphology of their typically coiled shell, exhibit a wide variety of dietary specializations (detritus feeders, suspension feeders, algal grazers, carnivores, parasites), and are adapted to very different marine, freshwater and terrestrial habitats worldwide (being particularly abundant in the tropics; Colgan et al., 2007). ...
... This latter group contains the majority of the diversity of caenogastropods, and was provisionally divided into three main groups, Littorinimorpha, Ptenoglossa, and Neogastropoda (=Stenoglossa) (Bouchet and Rocroi, 2005;Ponder and Lindberg, 1997). While the monophyly of the Caenogastropoda is generally not questioned either with morphology (Bouchet and Rocroi, 2005;Ponder and Lindberg, 1997;Strong, 2003; but see Haszprunar, 1988) or with molecules (Colgan et al., 2007;Harasewych et al., 1998;McArthur and Harasewych, 2003;Osca et al., 2014a;Zapata et al., 2014), the monophyly and phylogenetic relationships of the above mentioned main lineages within the group are under constant debate and revision (Colgan et al., 2007;Ponder et al., 2008). For instance, several morphological studies suggested that Architaenioglossa could represent a grade (Haszprunar, 1988;Ponder et al., 2008;Ponder and Lindberg, 1997) and considered Ptenoglossa non-monophyletic (Ponder and Lindberg, 1997), whereas several molecular studies fail to recover the monophyly of Littorinimorpha and Neogastropoda (Colgan et al., 2007;Cunha et al., 2009). ...
... While the monophyly of the Caenogastropoda is generally not questioned either with morphology (Bouchet and Rocroi, 2005;Ponder and Lindberg, 1997;Strong, 2003; but see Haszprunar, 1988) or with molecules (Colgan et al., 2007;Harasewych et al., 1998;McArthur and Harasewych, 2003;Osca et al., 2014a;Zapata et al., 2014), the monophyly and phylogenetic relationships of the above mentioned main lineages within the group are under constant debate and revision (Colgan et al., 2007;Ponder et al., 2008). For instance, several morphological studies suggested that Architaenioglossa could represent a grade (Haszprunar, 1988;Ponder et al., 2008;Ponder and Lindberg, 1997) and considered Ptenoglossa non-monophyletic (Ponder and Lindberg, 1997), whereas several molecular studies fail to recover the monophyly of Littorinimorpha and Neogastropoda (Colgan et al., 2007;Cunha et al., 2009). ...
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In order to further resolve the phylogenetic relationships within Caenogastropoda, the complete mitochondrial (mt) genomes of Cochlostoma hidalgoi (Cyclophoroidea), Naticarius hebraeus (Naticoidea), Galeodea echinophora (Tonnoidea), and Columbella adansoni (Buccinoidea), and the partial mt genome of Erosaria spurca (Cypraeoidea) were sequenced. All newly determined mt genomes conformed to the consensus gene order of caenogastropods, except that of C. hidalgoi, which differed in the relative positions of the trnD, trnQ, trnG, trnY, and trnT genes. Phylogenetic reconstruction of the caenogastropod tree was performed using probabilistic methods and based on the deduced amino acid sequences of concatenated mt protein coding genes. The reconstructed phylogeny recovered Architaenioglossa (superfamilies Cyclophoroidea, Ampullarioidea, and Viviparoidea) as a grade. The monophyly of Sorbeoconcha (all caenogastropods but Architaenioglossa) was supported by most but not all phylogenetic analyses (excluding Vermetoidea, which has a long branch). The relative phylogenetic position of Cerithioidea with respect to Hypsogastropoda remains unresolved. The monophyly of Hypsogastropoda (without Vermetoidea) is strongly supported. Within this clade, Littorinimorpha should be considered a grade. Several superfamilies (Abyssochrysoidea, Rissooidea, Truncatelloidea, and Naticoidea) branched off successively before a siphonate clade (Stromboidea, Cypraeoidea, Tonnoidea, Neogastropoda), which is strongly supported. The relative phylogenetic position of Vermetoidea could not be determined due to long-branch attraction artifacts. The superfamily Tonnoidea was recovered within Neogastropoda, which questions the monophyly of the latter as traditionally defined. We propose to resolve the polyphyly of Muricoidea by erecting two new superfamilies Volutoidea and Babylonioidea. Copyright © 2015 Elsevier Inc. All rights reserved.
... Mollusks are a diverse and abundant group, although often inconspicuous in the reef ecosystem. Among gastropods, species of Architectonicidae Gray, 1840, Coralliophilidae Chenu, 1859, Epitoniidae S. S. Berry, 1910, Muricidae Rafinesque, 1815, Nystiellidae Clench and Turner, 1952, Olividae Latreille, 1825, and Ovulidae Fleming, 1822, are known to live on stony and soft corals, hydroids, hydrocorals, discophores, siphonophores, gorgonians, zoanthids, and sea anemones, feeding on living cnidarian tissues [1][2][3][4][5]. ...
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A total of six genera and 10 species of marine gastropods belonging to the family Epitoniidae were collected from dredges of the continental slope off Brazil during the development of the REVIZEE (Live Resources of the Economic Exclusive Zone) Program. These species, referable to the genera Alora, Amaea, Cycloscala, Epitonium, Gregorioiscala, and Opalia, are reported from bathyal depths off northeastern Brazil. Alora sp., Gregorioiscala pimentai n. sp., and Opalia revizee n. sp. are species heretofore unknown to science. A list of the species of Epitonium and Opalia from the Atlantic coast of South America is presented based primarily on data from the literature. In addition, an overview of the biodiversity and distribution of the genera studied is presented for the Atlantic Ocean.
... This confusing pattern is partly driven by the lack of a clear phylogenetic (and thus systematic) framework for the entire Neogastropoda (Ponder et al., 2008). In fact, the use of morphological characters alone (either from the shell or from the anatomy of soft parts) for defining phylogenetic affinities is severely complicated, in neogastropods, by the strong tendency for parallel evolution of anatomical features. ...
... 13.16) and by combined morphological and molecular analyses (Ponder et al., 2008: 364 fig. 13.17), yet molecular analyses have often contradicted it (Ponder et al., 2008, and references therein). The tonnoideans (which often made neogastropods polyphyletic in several molecular phylogenies, as the velutinoidean Triviidae and Velutinidae sometimes did; e.g. ...
Article
The gastropod genus Colubraria includes marine shallow-water species from tropical, subtropical, and temperate rocky coral environments. At least six species are known to feed on fish blood. Although there is general consensus in placing Colubraria in the Neogastropoda, the actual relationships and the systematic position of Colubraria and related genera are unknown. This is partly the consequence of the lack of a clear phylogenetic framework for the Neogastropoda. This study attempts to propose a phylogenetic framework for the Neogastropoda, by testing: (1) a preliminary phylogenetic arrangement for a large number of recognized neogastropod families; (2) the position of Colubraria within the neogastropods; and (3) the relationships of Colubraria within one of the major neogastropod lineages. We used two different molecular data sets. The first set included representatives of at least 14 neogastropod families, for points (1) and (2), and was based on mitochondrial (16S, 12S, and cytochrome oxidase subunit I, COI) and nuclear (28S) DNA sequences, giving a total of 3443 aligned positions. The second data set, for point (3), included 30 buccinoid sequences from mitochondrial 16S, giving a total of 1029 aligned positions. We also studied the anatomy of the type species of Colubraria and compared it with other neogastropods within the new phylogenetic framework. The results included the first phylogeny of the neogastropod based on 50% of the recognized families. This clearly indicated that the nematoglossan Cancellariidae represent a basal offshoot of the monophyletic Neogastropoda, and that the toxoglossan Conoidea are the sister group to the Rachiglossa. Within the Rachiglossa, a colubrariid clade, worthy of family ranking, showed clear buccinoid affinities. Most of the anatomy of Colubraria is congruent with a buccinoid model. The peculiar anatomical features that do not conform to the buccinoid model seem to be related to the evolution of haematophagous feeding.