N. Puillandre, M. J. Tenorio

Journal of Molluscan Studies, Volume 83(2), May 2017, Pages 200–210

A question of rank: DNA sequences and radula characters reveal a new genus of cone snails (Gastropoda: Conidae)

doi: 10.1093/mollus/eyx011


Molecular phylogenies of cone snails have revealed that the c. 350 sequenced species are divided into four main lineages, Conus, Conasprella, Californiconus and Profundiconus. In a recent study, minute species (less than 8 mm) were for the first time included in a molecular phylogenetic tree and were shown to correspond to deep lineages, of similar status to the four previously recognized, and sister-group to Californiconus. They were attributed to the available generic names Lilliconus and Pseudolilliconus. In this article, we analyse, using morphological (shell and radula) and molecular characters (cox1 gene), several species of minute cone snails, and we conclude that the species considered as Pseudolilliconus in the previous study should actually be placed in a new genus, Pygmaeconus. By comparing the cox1 genetic distances calculated among the species of Lilliconus, Pygmaeconus and Californiconus with the genetic distances calculated among other cone snails species included in different subgenera and genera, and by comparing the estimated ages of Lilliconus and Pygmaeconus with the ages of other caenogastropod genera, we conclude that Lilliconus and Pygmaeconus can be considered at the genus rank.


S. Abalde, M. J. Tenorio, C. M. L. Afonso, R. Zardoya

Molecular Phylogenetics and Evolution Volume 112, April 2017, Pages 79-87

Mitogenomic phylogeny of cone snails endemic to Senegal

doi: 10.1016/j.ympev.2017.04.020


Cone snails attain in Senegal one of their highest peaks of species diversity throughout the continental coast of Western Africa. A total of 15 endemic species have been described, all placed in the genus Lautoconus. While there is ample data regarding the morphology of the shell and the radular tooth of these species, virtually nothing is known regarding the genetic diversity and phylogenetic relationships of one of the most endangered groups of cones. In this work, we determined the complete or near- complete (only lacking the control region) mitochondrial (mt) genomes of 17 specimens representing 11 endemic species (Lautoconus belairensis, Lautoconus bruguieresi, Lautoconus cacao, Lautoconus cloveri, Lautoconus cf. echinophilus, Lautoconus guinaicus, Lautoconus hybridus, Lautoconus senegalensis, Lautoconus mercator,Lautoconus taslei, and Lautoconus unifasciatus). We also sequenced the complete mt genome of Lautoconus guanche from the Canary Islands, which has been related to the cones endemic to Senegal. All mt genomes share the same gene arrangement, which conforms to the consensus reported for Conidae, Neogastropoda and Caenogastropoda. Phylogenetic analyses using probabilistic methods recovered three major lineages, whose divergence coincided in time with sea level and ocean current changes as well as temperature fluctuations during the Messinian salinity crisis and the Plio-Pleistocene transition. Furthermore, the three lineages corresponded to distinct types of radular tooth (robust, small, and elongated), suggesting that dietary specialization could be an additional evolutionary driver in the diversification of the cones endemic to Senegal. The reconstructed phylogeny showed several cases of phenotypic convergence (cryptic species) and questions the validity of some species (ecotypes or phenotypic plasticity), both results having important taxonomic and conservation consequences.

J. E. Uribe, N. Puillandre, R. Zardoya

Molecular Phylogenetics and Evolution Volume 107, February 2017, Pages 142-151

Beyond Conus: Phylogenetic relationships of Conidae based on complete mitochondrial genomes

doi: 10.1016/j.ympev.2016.10.008


Understanding how the extraordinary taxonomic and ecological diversity of cone snails (Caenogastropoda: Conidae) evolved requires a statistically robust phylogenetic framework, which thus far is not available. While recent molecular phylogenies have been able to distinguish several deep lineages within the family Conidae, including the genera Profundiconus, Californiconus, Conasprella, and Conus (and within this one, several subgenera), phylogenetic relationships among these genera remain elusive. Moreover, the possibility that additional deep lineages may exist within the family is open. Here, we reconstructed with probabilistic methods a molecular phylogeny of Conidae using the newly sequenced complete or nearly complete mitochondrial (mt) genomes of the following nine species that represent all main Conidae lineages and potentially new ones: Profundiconus teramachii, Californiconus californicus, Conasprella wakayamaensis, Lilliconus sagei, Pseudolilliconus traillii, Conus (Kalloconus) venulatus, Conus (Lautoconus) ventricosus, Conus (Lautoconus) hybridus, and Conus (Eugeniconus) nobilis. To test the monophyly of the family, we also sequenced the nearly complete mt genomes of the following three species representing closely related conoidean families: Benthomangelia sp. (Mangeliidae), Tomopleura sp. (Borsoniidae), and Glyphostoma sp. (Clathurellidae). All newly sequenced conoidean mt genomes shared a relatively constant gene order with rearrangements limited to tRNA genes. The reconstructed phylogeny recovered with high statistical support the monophyly of Conidae and phylogenetic relationships within the family. The genus Profundiconus was placed as sister to the remaining genera. Within these, a clade including Californiconus and Lilliconus + Pseudolilliconus was the sister group of Conasprella to the exclusion of Conus. The phylogeny included a new lineage whose relative phylogenetic position was unknown (Lilliconus) and uncovered thus far hidden diversity within the family (Pseudolilliconus). Moreover, reconstructed phylogenetic relationships allowed inferring that the peculiar diet of Californiconus based on worms, mollusks, crustaceans and fish is derived, and reinforce the hypothesis that the ancestor of Conidae was a worm hunter. A chronogram was reconstructed under an uncorrelated relaxed molecular clock, which dated the origin of the family shortly after the Cretaceous-Tertiary boundary (about 59 million years ago) and the divergence among main lineages during the Paleocene and the Eocene (56-30 million years ago).

S. Abalde, M. J. Tenorio, C. M. L. Afonso, Juan E. Uribe, Ana M. Echeverry, R. Zardoya

BMC Evolutionary Biology Volume 17(1), November 2017, Article 231
Phylogenetic relationships of cone snails endemic to Cabo Verde based on mitochondrial genomes
doi: 10.1186/s128620171069x


Background: Due to their great species and ecological diversity as well as their capacity to produce hundreds of different toxins, cone snails are of interest to evolutionary biologists, pharmacologists and amateur naturalists alike. Taxonomic identification of cone snails still relies mostly on the shape, color, and banding patterns of the shell. However, these phenotypic traits are prone to homoplasy. Therefore, the consistent use of genetic data for species delimitation and phylogenetic inference in this apparently hyperdiverse group is largely wanting. Here, we reconstruct the phylogeny of the cones endemic to Cabo Verde archipelago, a well known radiation of the group, using complete mitochondrial (mt) genomes.

Results: The reconstructed phylogeny grouped the analyzed species into two main clades, one including Kalloconus from West Africa sister to Trovaoconus from Cabo Verde and the other with a paraphyletic Lautoconus due to the sister group relationship of Africonus from Cabo Verde and Lautoconus ventricosus from Mediterranean Sea and neighboring Atlantic Ocean to the exclusion of Lautoconus endemic to Senegal (plus Lautoconus guanche from Mauritania, Morocco, and Canary Islands). Within Trovaoconus, up to three main lineages could be distinguished. The clade of Africonus included four main lineages (named I to IV), each further subdivided into two monophyletic groups. The reconstructed phylogeny allowed inferring the evolution of the radula in the studied lineages as well as biogeographic patterns. The number of cone species endemic to Cabo Verde was revised under the light of sequence divergence data and the inferred phylogenetic relationships.

Conclusions: The sequence divergence between continental members of the genus Kalloconus and island endemics ascribed to the genus Trovaoconus is low, prompting for synonymization of the latter. The genus Lautoconus is paraphyletic. Lautoconus ventricosus is the closest living sister group of genus Africonus. Diversification of Africonus was in allopatry due to the direct development nature of their larvae and mainly triggered by eustatic sea level changes during the Miocene-Pliocene. Our study confirms the diversity of cone endemic to Cabo Verde but significantly reduces the number of valid species. Applying a sequence divergence threshold, the number of valid species within the sampled Africonus is reduced to half.

E. Monnier, M. J. Tenorio

Xenophora Taxonomy, Volume 6, January 2015, Pages 47–51

Malagasyconus (Gastropoda: Conidae), a new genus from Southern Madagascar


The taxon Malagasyconus new genus is introduced. This genus contains two living species: Malagasyconus lozeti (Richard, 1980) and M. bonfigliolii (Bozzetti, 2010), which have been collected in deep water in Southern Madagascar. Malagasyconus n. gen. is compared to the genera Plicaustraconus Moolenbeek, 2008 and Quasiconus Tucker & Tenorio, 2009.

N. Puillandre, T.F. Duda Jr., B.M. Olivera, C. Meyer, P. Bouchet

Journal of Molluscan Studies, Volume 81(1), January 2015, Pages 1–23

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

 doi: 10.1093/mollus/eyu055   

 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 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 confidence 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.



N. Puillandre, P. Bouchet, T.F. Duda Jr., S. Kauferstein, A.J. Kohn, B.M. Olivera, M. Watkins, C. Meyer

Molecular Phylogenetics and Evolution, Volume 78, September 2014, Pages 290–303

Molecular phylogeny and evolution of the cone snails (Gastropoda, Conoidea)


 We present a large-scale molecular phylogeny that includes 320 of the 761 recognized valid species of the cone snails (Conus), one of the most diverse groups of marine molluscs, based on three mitochondrial genes (COI, 16S rDNA and 12S rDNA). This is the first phylogeny of the taxon to employ concatenated sequences of several genes, and it includes more than twice as many species as the last published molecular phylogeny of the entire group nearly a decade ago. Most of the numerous molecular phylogenies published during the last 15 years are limited to rather small fractions of its species diversity. Bayesian and maximum likelihood analyses are mostly congruent and confirm the presence of three previously reported highly divergent lineages among cone snails, and one identified here using molecular data. About 85% of the species cluster in the single Large Major Clade; the others are divided between the Small Major Clade (∼12%), the Conus californicus lineage (one species), and a newly defined clade (∼3%). We also define several subclades within the Large and Small major clades, but most of their relationships remain poorly supported. To illustrate the usefulness of molecular phylogenies in addressing specific evolutionary questions, we analyse the evolution of the diet, the biogeography and the toxins of cone snails. All cone snails whose feeding biology is known inject venom into large prey animals and swallow them whole. Predation on polychaete worms is inferred as the ancestral state, and diet shifts to molluscs and fishes occurred rarely. The ancestor of cone snails probably originated from the Indo-Pacific; rather few colonisations of other biogeographic provinces have probably occurred. A new classification of the Conidae, based on the molecular phylogeny, is published in an accompanying paper.

New Advances in Taxonomy

New genera, subgenera, and relevant information on cone Taxonomy reported after the publication of the Illustrated Catalog of the Living Cone Shells

In this section we will summarize all the information on the new genera, subgenera, changes in generic ascription for species, and in general all relevant information related to cone Taxonomy, including digests of important papers appeared on Conoidean phylogeny.