Taxonomic revision of West African cone snails (Gastropoda: Conidae) based upon mitogenomic studies: implications for conservation

  • Manuel J. Tenorio Departamento CMIM y Química Inorgánica – Instituto de Biomoléculas (INBIO), Facultad de Ciencias, Torre Norte, 1ª Planta, Universidad de Cadiz, 11510 Puerto Real, Cadiz
  • Samuel Abalde Museo Nacional de Ciencias Naturales (MNCN-CSIC), José Gutiérrez Abascal 2, 28006 Madrid
  • José R. Pardos-Blas Museo Nacional de Ciencias Naturales (MNCN-CSIC), José Gutiérrez Abascal 2, 28006 Madrid
  • Rafael Zardoya Museo Nacional de Ciencias Naturales (MNCN-CSIC), José Gutiérrez Abascal 2, 28006 Madrid
Keywords: mitochondrial genome, phylogeny, Conidae, West Africa, conservation

Abstract

In the last few years, a sharp increase in the number of descriptions of new species of West African cone snails, particularly from the Cabo Verde Archipelago, has taken place. In previous studies, we used mitogenome sequences for reconstructing robust phylogenies, which comprised in total 120 individuals representing the majority of species (69.7%) described from this biogeographical region (except Angolan endemics) and grouped into seven genera within the family Conidae. Here, we add another 12 individuals representing endemic species that were missing in the previous studies. We use the phylogenies to identify monophyletic groups and a genetic divergence threshold (0.2% uncorrected p distance) to determine the number of valid species. As a result, the number of valid West African cone species could be drastically reduced to at least 40%, indicating that some recent poor-quality descriptions loosely based on phenotypic characters prone to convergence such as the shape and color patterns of the shell have contributed substantially to taxonomic inflation. Several previously accepted species with a reduced geographical distribution now become phenotypic forms of the remaining valid species, which increase their distribution ranges. In contrast, several cryptic species are now uncovered and described. For instance, Africonus insulae sp. nov. and Kalloconus canariensis sp. nov. are hereby introduced as new species. A detailed systematic account with illustrations and relevant information is presented. Lectotypes are designated for Conus trochulus and Conus irregularis, and neotypes for Conus crotchii and Conus diminutus. According to our results, it is strongly recommended that any future introduction of new taxa names for cone snails from West Africa should be supported by molecular and/or anatomical rather than exclusively shell morphological data. The taxonomic decisions here taken have direct implications for conservation and will eventually require re-evaluation of the Red List risk status of an important number of species.

References

Abalde S., Tenorio M.J., Afonso C.M.L., Uribe J.E., Echeverry A.M. & Zardoya R. 2017a. Phylogenetic relationships of cone snails endemic to Cabo Verde based on mitochondrial genomes. BMC Evolutionary Biology 17: 231. https://doi.org/10.1186/s12862-017-1069-x

Abalde S., Tenorio M.J., Afonso C.M.L. & Zardoya R. 2017b. Mitogenomic phylogeny of cone snails endemic to Senegal. Molecular Phylogenetics and Evolution 112: 79–87. https://doi.org/10.1016/j.ympev.2017.04.020

Abalde S., Tenorio M.J., Afonso C.M.L. & Zardoya R. 2018. Conotoxin diversity in Chelyconus ermineus (Born, 1778) and the convergent origin of piscivory in the Atlantic and Indo-Pacific cones. Genome Biology and Evolution 10 (10): 2643–2662. https://doi.org/10.1093/gbe/evy150

Abalde S., Tenorio M.J., Uribe J.E. & Zardoya R. 2019. Conidae phylogenomics and evolution. Zoologica Scripta 48: 194–214. https://doi.org/10.1111/zsc.12329

Abalde S., Tenorio M.J., Afonso C.M.L. & Zardoya R. 2020. Comparative transcriptomics of the venoms of continental and insular radiations of West African cone snails. In press. https://doi.org/10.1098/rspb.2020.0794

Adams A. 1854. Descriptions of new species of the genus Conus, from the collection of Hugh Cuming, Esq. Proceedings of the Zoological Society of London 1854: 116–119. Available from http://biology.burke.washington.edu/conus/catalogue/sources/Adams,%20A.,%201854.%20Proc.%20Zool.%20Soc.%20Lond..pdf [accessed 29 Mar. 2020].

Afonso C.M.L. & Tenorio M.J. 2004. Description of a new offshore species of Conus from the Cape Verde Archipelago (Gastropoda, Conidae). La Conchiglia: International Shell Magazine 36 (310): 33–40.

Afonso C.M.L. & Tenorio M.J. 2011. A new, distinct endemic Africonus species (Gastropoda, Conidae) from São Vicente Island, Cape Verde Archipelago, West Africa. Gloria Maris 50 (5):125–136. Available from https://storage.googleapis.com/conchology-general-images/images/gloriamaris/pdfs/2011/Vol50-5.pdf [accessed 23 May 2020]

Afonso C.M.L. & Tenorio M.J. 2014. Recent findings from the islands of Maio and Boa Vista in the Cape Verde Archipelago, West Africa: description of three new Africonus species (Gastropoda: Conidae). Xenophora Taxonomy 3: 47–60.

Bandel K. & Wils E. 1977. On Conus mediterraneus and Conus guinaicus. Basteria 41: 33–45. Available from http://natuurtijdschriften.nl/download?type=document;docid=596671 [accessed 29 Mar. 2020].

Barros e Cunha J.G. 1933. Catálogo descritivo das conchas exóticas da colecção António Augusto de Carvalho Monteiro Família Conidae. Memórias e Estudos do Museu Zoológico da Universidade de Coimbra sér. 1 71: 1–224.

Bettocchi M. 2018. Genuanoconus genuanus (Linnaeus, 1758): a species very variable in pattern and profile. The Cone Collector 31: 6–36. Available from http://theconecollector.com/lib/docs/tcc_31.pdf.zip [accessed 29 Mar. 2020].

Boivin A. 1864. Description de cinq espèces nouvelles du genre Conus. Journal de Conchyliologie 12: 33–40.

Borges L.M.S., Hollatz C., Lobo J., Cunha A.M., Vilela A.P., Calado G., Coelho R., Costa A.C., Ferreira M.S.G., Costa M.H. & Costa F.O. 2016. With a little help from DNA barcoding: investigating the diversity of Gastropoda from the Portuguese coast. Scientific Reports 6: 20226–20226. https://doi.org/10.1038/srep20226

Born I. von 1778. Index Rerum Naturalium Musei Caesarei Vindobonensis. Pars Prima, Testacea. Available from http://biology.burke.washington.edu/conus/catalogue/sources/Born,%20I.,%201778.%20%20Index%20Rerum%20Naturalium.pdf [accessed 29 Mar. 2020].

Boyer F. & Pelorce J. 2009. Description d’un nouveau Conus (Gastropoda: Conidae) du Sénégal dans le groupe Conus mediterraneus. Novapex 10 (1): 25–32. Available from https://www.biodiversitylibrary.org/page/42270880 [accessed 29 Mar. 2020].

Burnay L.P. & Monteiro A.A. 1977. Seashells from Cape Verde Islands. Published by the authors, Lisbon.

Bruguière J.G. 1792. Cone. In: Encyclopédie méthodique: Histoire naturelle des Vers 1: 586–757. Panckouke, Paris. https://doi.org/10.5962/bhl.title.49857

Chernomor O., von Haeseler A. & Minh B.Q. 2016. Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology 65: 997–1008. https://doi.org/10.1093/sysbio/syw037

Clench W.J. 1942. The genus Conus in the western Atlantic. Johnsonia 1 (6): 1–40.

Coomans H.E., Moolenbeek R.G. & Wils E. 1980. Alphabetical revision of the (sub)species in recent Conidae 3. albus to antillarum with the description of Conus algoensis agulhasi, nov. subspecies. Basteria 44 (1/4): 17–49. Available from http://natuurtijdschriften.nl/download?type=document&docid=596716 [accessed 29 Mar. 2020].

Coomans H.E., Moolenbeek R.G. & Wils E. 1981. Alphabetical revision of the (sub)species in recent Conidae 4. aphrodite to azona with the description of Conus arenatus bizona nov. subspecies. Basteria 45 (1/3): 3–55. Available from http://natuurtijdschriften.nl/download?type=document&docid=596730 [accessed 29 Mar. 2020].

Coomans H.E., Moolenbeek R.G. & Wils E. 1982. Alphabetical revision of the (sub)species in recent Conidae 5. baccatus to byssinus, including Conus brettinghami nomen novum. Basteria 46 (1/4): 3–67. Available from http://natuurtijdschriften.nl/download?type=document&docid=596751 [accessed 29 Mar. 2020].

Coomans H.E., Moolenbeek R.G. & Wils E. 1985a. Alphabetical revision of the (sub)species in recent Conidae 7. cingulatus to cylindraceus, including Conus shikamai nomen novum. Basteria 48 (6): 223–311. Available from http://natuurtijdschriften.nl/download?type=document&docid=596791 [accessed 29 Mar. 2020].

Coomans H.E., Moolenbeek R.G. & Wils E. 1985b. Alphabetical revision of the (sub)species in recent Conidae 8. dactylosus to dux. Basteria 49 (4/6): 145–196. Available from http://natuurtijdschriften.nl/download?type=document&docid=596813 [accessed 29 Mar. 2020].

Coomans H.E., Moolenbeek R.G. & Wils E. 1986. Alphabetical revision of the (sub)species in recent Conidae 9. ebraeus to extraordinarius with the description of Conus elegans ramalhoi nov. subspecies. Basteria 50 (4/6): 93–150. Available from http://natuurtijdschriften.nl/download?type=document&docid=596828 [accessed 7 Apr. 2020].

Cossignani T. 2014. Dieci nuovi coni da Capo Verde. Malacologia Mostra Mondiale 82 (1): 18–29.

Cossignani T. & Allary A. 2019. Lautoconus (Lautoconus) stimpsonorum: nuova specie dal Senegal. Malacologia Mostra Mondiale 104: 21-23.

Cossignani T. & Fiadeiro R. 2014a. Quattro nuovi coni da Capo Verde. Malacologia Mostra Mondiale 83 (2): 14–19.

Cossignani T. & Fiadeiro R. 2014b. Cinque nuovi coni da Capo Verde. Malacologia Mostra Mondiale 84 (3): 21–27.

Cossignani T. & Fiadeiro R. 2015a. Tre nuovi coni da Capo Verde. Malacologia Mostra Mondiale 86: 17–21.

Cossignani T. & Fiadeiro R. 2015b. Due Nuovi Coni da Capo Verde. Malacologia Mostra Mondiale 87: 3–5.

Cossignani T. & Fiadeiro R. 2015c. Due Nuovi Coni da Capo Verde. Malacologia Mostra Mondiale 88: 3–5.

Cossignani T. & Fiadeiro R. 2017a. Otto nuovi coni da Capo Verde. Malacologia Mostra Mondiale 94: 26–36.

Cossignani T. & Fiadeiro R. 2017b. Un nuovo cono da Capo Verde. Malacologia Mostra Mondiale 96: 32–34.

Cossignani T. & Fiadeiro R. 2017c. Un nuovo cono da Capo Verde. Malacologia Mostra Mondiale 97: 52–53.

Cossignani T. & Fiadeiro R. 2018a. Quattro nuovi coni da Capo Verde. Malacologia Mostra Mondiale 98: 14–20.

Cossignani T. & Fiadeiro R. 2018b. Africonus perrineae: un nuovo cono da Capo Verde. Malacologia Mostra Mondiale 99: 17–19.

Cossignani T. & Fiadeiro R. 2018c. Africonus tarafensis: un nuovo cono da Capo Verde. Malacologia Mostra Mondiale 101: 32–34.

Cossignani T. & Fiadeiro R. 2019a. Kalloconus (Trovaoconus) josefiadeiroi un nuovo cono da Capo Verde. Malacologia Mostra Mondiale 102: 30–32.

Cossignani T. & Fiadeiro R. 2019b. Un nuovo cono da Capo Verde. Malacologia Mostra Mondiale 104: 16–17.

Cunha R.L., Castilho R., Rüber L. & Zardoya R. 2005. Patterns of cladogenesis in the venomous marine gastropod genus Conus from the Cabo Verde Archipelago. Systematic Biology 54: 634–650. https://doi.org/10.1080/106351591007471

Cunha R.L., Tenorio M.J., Afonso C., Castilho R. & Zardoya R. 2008. Replaying the tape: recurring biogeographical patterns in Cabo Verde Conus after 12 million years. Molecular Ecology 17: 885–901. https://doi.org/10.1111/j.1365-294X.2007.03618.x

Cunha R.L., Grande C. & Zardoya R. 2009. Neogastropod phylogenetic relationships based on entire mitochondrial genomes. BMC Evolutionary Biology 9: 210. https://doi.org/10.1186/1471-2148-9-210

Cunha R.L., Lima F.P., Tenorio M.J., Ramos A.A., Castilho R. & Williams S.T. 2014. Evolution at a different pace: distinctive phylogenetic patterns of cone snails from two ancient oceanic archipelagos. Systematic Biology 63: 971–987. https://doi.org/10.1093/sysbio/syu059

da Motta A.J. 1982. Seventeen new cone shell names (Gastropoda: Conidae). Publicações Ocasionais da Sociedade Portuguesa de Malacologia 1: 1–20.

da Motta A.J. 1991. A Systematic Classification of the Gastropod Family Conidae at the Generic Level. La Conchiglia, Rome.

Dillwyn L.W. 1817. A Descriptive Catalogue of Recent Shells, Arranged According to the Linnean Method; with Particular Attention to the Synonymy, vol. 1. J. and A. Arch, London. Available from http://biology.burke.washington.edu/conus/catalogue/sources/Dillwyn,%20L.W.,%201817.%20Descr.%20Cat.%20Rec.%20Shells,%20i..pdf [accessed 29 Mar. 2020].

Duda T.F. & Rolán E. 2005. Explosive radiation of Cabo Verde Conus, a marine species flock. Molecular Ecology 14: 267–272. https://doi.org/10.1111/j.1365-294X.2004.02397.x

Duda T.F., Kohn A.J. & Palumbi S.R. 2001. Origins of diverse feeding ecologies within Conus, a genus of venomous marine gastropods. Biological Journal of the Linnean Society 73: 391–409. https://doi.org/10.1006/bijl.2001.0544

Echterbille J., Gilles N., Araóz R., Mourier G., Amar M., Servent D., De Pauw E. & Quinton L. 2017. Discovery and characterization of EIIB, a new -conotoxin from Conus ermineus venom by nAChRs affinity capture monitored by MALDI-TOF/TOF mass spectrometry. Toxicon 130: 1–10. https://doi.org/10.1016/j.toxicon.2017.02.023

Elsen J. 1983. Taxonomy of some Cape Verde Conidae with description of a novel species. Informations de la Société belge de Malacologie 11: 180–189.

Faber M. 2011. The holy grail of Louis Charles Kiener’s “Spécies général des coquilles vivantes”. Miscellanea Malacologica 5 (3): 61–70. Available from http://olivirv.myspecies.info/sites/olivirv.myspecies.info/files/The%20holy%20grail%20of%20Louis%20Charles%20-%20Faber%2C%20M.J_.pdf [accessed 29 Mar. 2020].

Ferrario M. 1983. La Piu’ Grande Enciclopedia delle Conchiglie di Tutto il Mondo. Alberto Peruzzi, Milan.

Filmer R.M. 2011. Nomenclature and Taxonomy in Living Conidae, 1st Ed. Available from http://www.theconecollector.com/filmer/index.html [accessed 29 Mar. 2020].

Fischer von Waldheim G. 1807. Museum - Demidoff, ou Catalogue systématique et raisonné des Curiosités de la Nature et de l’Art, données à l’Université impériale de Moscou par son Excellence Monsieur Paul de Demidoff. Tome Troisième. Moscow. https://doi.org/10.5962/bhl.title.160942

Folmer O., Black M., Hoeh W., Lutz R. & Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294–299.

Freitas R., Romeiras M., Silva L., Cordeiro R., Madeira P., González J.A., Wirtz P., Falcón J.M., Brito A., Floeter S.R., Afonso P., Porteiro F., Viera-Rodríguez M.A., Neto A.I., Haroun R., Farminhão J.N.M., Rebelo A.C., Baptista L., Melo C.S., Martínez A., Núñez J., Berning B., Johnson M.E. & Ávila S.P. 2019. Restructuring of the “Macaronesia” biogeographic unit: a marine multi-taxon biogeographical approach. Scientific Reports 9: 15792. https://doi.org/10.1038/s41598-019-51786-6

Gmelin J.F. 1791. Caroli a Linné. Systema Naturae per Regna Tria Naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, 13th Ed. Vol. 1. Impensis Georg. Emanuel. Beer, Leipzig [Lipsiae]. https://doi.org/10.5962/bhl.title.36932

Grande C., Templado J. & Zardoya R. 2008. Evolution of gastropod mitochondrial genome arrangements. BMC Evolutionary Biology 8: 61. https://doi.org/10.1186/1471-2148-8-61

Gulden G.J., Moolenbeek R.G. & Goud J. 2017. A new Lautoconus (Gastropoda: Conidae) from Senegal. Miscellanea Malacologica 7 (3): 49‒53.

Kiener L.C. 1845–1859. Genre Cone. (Conus, Lin.) Spécies général et iconographie des coquilles vivantes comprenant la collection du Muséum d’Histoire naturelle de Paris, collection Lamarck, celle du Prince Masséna et les découverts récente des voyageurs. Rousseau, Paris [pp. 1–64, 1846; 65–176, 1847; 177–224, 1848; 225–272, 1859; 273–379, 1849–50; plates 1–111, 1845]. https://doi.org/10.5962/bhl.title.39135

Kohn A.J. 1963. Type specimens and identity of the described species of Conus. I. The species described by Linnaeus, 1758–1767. Journal of the Linnean Society of London, Zoology 44 (302): 740–768. https://doi.org/10.1111/j.1096-3642.1963.tb01641.x

Kohn A.J. 1964. Type specimens and identity of the described species of Conus. II. The species described by Solander, Chemnitz, Born, and Lightfoot between 1766 and 1786. Journal of the Linnean Society of London, Zoology 45 (304): 151–167. https://doi.org/10.1111/j.1096-3642.1964.tb00492.x

Kohn A.J. 1968. Type specimens and identity of the described species of Conus. IV: the species described by Hwass, Bruguière and Olivi in 1792. Journal of the Linnean Society of London, Zoology 47 (313): 431–503. https://doi.org/10.1111/j.1096-3642.1968.tb00550b.x

Kohn A.J. 1975. Type specimens and identity of the described species of Conus. V. The species described by Salis Marschlins and Röding, 1793–1798. Zoological Journal of the Linnean Society 57: 185–226. https://doi.org/10.1111/j.1096-3642.1975.tb00816.x

Kohn A.J. 1992. A Chronological Taxonomy of Conus, 1758–1840. Smithsonian Institution Press, Washington D.C., USA.

Kohn A.J., Nishi M. & Pernet B. 1999. Snail spears and scimitars: a character analysis of Conus radular teeth. Journal of Molluscan Studies 65: 461–481. https://doi.org/10.1093/mollus/65.4.461

Küster H.C. 1838. Coneae, oder Conidae, I. Conus, Linne. Systematisches Conchylien-Cabinet von Martini und Chemnitz, neu herausgegeben und vervollständigt von H.C. Küster. Band 4, Abtheilung 2. Raspe, Nürnberg.

Lamarck J.B.P. 1810. Sur la détermination des espèces parmi les animaux sans vertèbres, et particulièrement parmi les Mollusques testacés. Annales du Muséum d’histoire naturelle 15: 1–442. Available from https://www.biodiversitylibrary.org/item/44182#page/8/mode/1up [accessed 7 Apr. 2020].

Lanfear R., Frandsen P.B., Wright A.M., Senfeld T. & Calcott B. 2017. PartitionFinder2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution 34 (3): 772–3. https://doi.org/10.1093/molbev/msw260

Lauer J.M. 1993a. Description of a new species and a new subspecies of Conus (Mollusca: Prosobranchia: Conidae) from the Canary Islands. Apex 8 (1/2): 37–50. Available from http://www.vliz.be/imisdocs/publications/294939.pdf [accessed 29 Mar. 2020].

Lauer J.M. 1993b. Conus cacao Ferrario, 1983, taxonomical and systematic context (Mollusca: Prosobranchia: Conidae). Apex 8(3): 115–127. Available from http://www.vliz.be/imisdocs/publications/294973.pdf [accessed 29 Mar. 2020].

Lightfoot J. 1786. A Catalogue of the Portland Museum, lately the Property of the Duchefs Dowager of Portland, deceased; which will be sold by auction by Mr. Skinner & Co. London. Available from http://resolver.sub.uni-goettingen.de/purl?PPN603451020 [accessed 29 Mar. 2020].

Link H.F. 1807. Beschreibung der Naturalien-Sammlung der Universität zu Rostock, Mollusken. Part 3: 101–160. Available from http://biology.burke.washington.edu/conus/catalogue/sources/Link,%201807.%20Besch.%20Nat.-Samml.%20Univ.%20Rostock.%203e%20Abth...pdf [accessed 29 Mar. 2020].

Linnaeus C. 1758. Systema Naturae per regna tria naturae, secundum classes ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio Decima, Reformata. Laurentii Salvii, Stockholm [Holmiae]. https://doi.org/10.5962/bhl.title.542

Linnaeus C. 1767. Systema Naturae per regna tria naturae, secundum classes ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio Duodecima, Reformata. Laurentii Salvii, Stockholm [Holmiae]. https://doi.org/10.5962/bhl.title.68927

Lister M., Lister S. & Lister A. 1688. Historiæ [sive Synopsis methodicæ] Conchyliorum [quorum omnium picturæ, ad vivum delineatæ, exhibētur.] Liber IV. qui est de buccinis marinis etiam vermiculj, dentalia, et patellæ numerantur. ibidem. London. Available from http://resolver.sub.uni-goettingen.de/purl?PPN470145102 [accessed 23 May 2020]

Martini F.H.W. 1773. Neues Systematisches Conchylien-Cabinet, 2. Nürnberg. Available from https://doi.org/10.5962/bhl.title.43760 [accessed 23 May 2020]

Miller M.A., Pfeiffer W. & Schwartz T. 2010. Creating the CIPRES science gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE): 1–8. 14 Nov. 2010, New Orleans, LA.

Monnier E. & Limpalaër L. 2010. Conus dorotheae (Gastropoda, Conidae) a new species of cone from the Cape Verde Peninsula in Senegal. Visaya 2 (6): 73–80.

Monnier E., Limpalaër L., Robin A. & Roux C. 2018. A Taxonomic Iconography of Living Conidae, Vols 1 and 2. ConchBooks, Harxheim.

Monteiro A., Fernandes C. & Rolán E. 1995. A new species in the genus Conus from Boavista Island, Cape Verde Islands. World Shells 12: 8–13.

Monteiro A., Tenorio M.J. & Poppe G.T. 2004. A Conchological Iconography; The Family Conidae: the West African and Mediterranean Species of Conus. ConchBooks, Hackenheim.

Monteiro A., Afonso C.M.L. & Rosa G.D. 2014. Designation of a neotype of Africonus maioensis (Trovao, Rolán & Felix-Alves, 1990). Xenophora Taxonomy 4: 3.

Neves J.L.B., Lin Z., Imperial J.S., Antunes A., Vasconcelos V., Olivera B. & Schmidt E.W. 2015. Small molecules in the cone snail arsenal. Organic Letters 17 (20): 4933–4935. https://doi.org/10.1021/acs.orglett.5b02389

Nguyen L.T., Schmidt H.A., von Haeseler A. & Minh B.Q. 2015. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32: 268–274. https://doi.org/10.1093/molbev/msu300

Nolf F. & Verstraeten J. 2008. Conus trencarti (Mollusca: Gastropoda: Conidae): a new cone from Senegal. Neptunea 7 (4): 1–3.

Osca D., Templado J. & Zardoya R. 2015. Caenogastropod mitogenomics. Molecular Phylogenetics and Evolution 93: 118–128. https://doi.org/10.1016/j.ympev.2015.07.011.

Palumbi S.R., Martin A., Romano S., McMillan W.O., Stice L. & Grabowski G. 1991. The Simple Fool's Guide to PCR. Ver. 2. University of Hawaii. Honolulu.

Perry G. 1811. Conchology, or The Natural History of Shells; Containing a New Arrangement of the Genera and Species. William Miller, London. https://doi.org/10.5962/bhl.title.64363

Peters H., O’Leary B.C., Hawkins J.P., Carpenter K.E. & Roberts C.M. 2013. Conus: first comprehensive conservation Red List assessment of a marine gastropod mollusc genus. PLoS ONE 8: e83353. https://doi.org/10.1371/journal.pone.0083353

Peters H., O’Leary B.C., Hawkins J.P. & Roberts C.M. 2016. The cone snails of Cabo Verde: marine endemism at a terrestrial scale. Global Ecology and Conservation 7: 201–213. https://doi.org/10.1016/j.gecco.2016.06.006

Petuch E.J. 1975a. A review of the small Mauretanian cone shells with the description of a new genus and a new species (Gastropoda: Conidae). The Veliger 17 (3): 262–263.

Petuch E.J. 1975b. Two new cone species from Senegal, West Africa. The Veliger 18 (2): 180–182.

Petuch E.J. & Berschauer D.P. 2016. A new species of cone shell (Gastropoda: Conidae) from the Saharan coast of northwestern Africa. The Festivus 48 (2): 93–99.

Petuch E.J. & Berschauer D.P. 2018. A new Lautoconus species radiation from Gambia, West Africa. The Festivus 50 (3): 164–172.

Phuong M.A., Alfaro M.E., Mahardika G.N., Marwoto R.M., Prabowo R.E., von Rintelen T., Vogt P.W.H., Hendricks J.R. & Puillandre N. 2019. Lack of signal for the impact of conotoxin gene diversity on speciation rates in cone snails. Systematic Biology 68 (5): 781–796. https://doi.org/10.1093/sysbio/syz016

Pin M. & Leung Tack K.D. in Pin M. 1989a. Une nouvelle espèce de gastéropode prosobranche des côtes du Sénégal: Conus belairensis n. sp. Publicações Ocasionais da Sociedade Portuguesa de Malacologia 14: 33–38.

Pin M. & Leung Tack K.D. in Pin M. 1989b. Étude des canaux anaux des Conidae du Sénégal et leur importance dans la détermination des espèces. Publicações Ocasionais da Sociedade Portuguesa de Malacologia 14: 49–78.

Pin M. & Leung Tack K.D. 1995. The cones of Senegal. La Conchiglia: International Shell Magazine 277: 1–56.

Puillandre N., Bouchet P., Duda T.F., Kauferstein S., Kohn A.J., Olivera B.M. & Meyer C. 2014. Molecular phylogeny and evolution of the cone snails (Gastropoda, Conoidea). Molecular Phylogenetics and Evolution 78: 290–303. https://doi.org/10.1016/j.ympev.2014.05.023

Puillandre N., Duda T.F., Meyer C.P., Olivera B.M. & Bouchet P. 2015. One, four or 100 genera? A new classification of the cone snails. Journal of Molluscan Studies 81: 1–23. https://doi.org/10.1093/mollus/eyu055

Quinton L., Servent D., Girard E., Molgó J., Le Caer J.P., Malosse C., Haidar el A., Lecoq A., Gilles N. & Chamot-Rooke J. 2013. Identification and functional characterization of a novel α-conotoxin (EIIA) from Conus ermineus. Analytical and Bioanalytical Chemistry 405 (15): 5341–5351. https://doi.org/10.1007/s00216-013-6926-x

Raybaudi G. 1992. Preliminary studies for a revision of subgenus Lautoconus (Monterosato,1923) in the Mediterranean sea. Part I: on the true identity of Conus desidiosus A. Adams, 1854. La Conchiglia: International Shell Magazine 23 (263): 10–15.

Reeve L.A. 1843–44. Monograph of the Genus Conus. Conchologia Iconica, I: Figures and Descriptions of the Shells of Molluscs; with Remarks on their Affinities, Synonymy, and Geographical Distribution, 1. Conus. Available from http://biology.burke.washington.edu/conus/catalogue/sources/Reeve,%201843-9.%20Conch.%20Icon..%20i.%20Conus..pdf [accessed 29 Mar. 2020].

Reeve L.A. 1848–49. Conus. Emendations applicable to the Monographs of the Conchologia Iconica, and Iconographie des Coquilles Vivantes, Vol. 1. Reeve Brothers, London. Available from http://biology.burke.washington.edu/conus/catalogue/sources/Reeve,%201848-9.%20Conch.%20Icon..%20i.%20Conus.%20Suppl..pdf [accessed 29 Mar. 2020].

Rivera-Ortiz J.A., Cano H. & Marí F. 2011. Intraspecies variability and conopeptide profiling of the injected venom of Conus ermineus. Peptides 32 (2): 306–316. https://doi.org/10.1016/j.peptides.2010.11.014

Röckel D. 1978. The rediscovery of Conus siamensis. Hawaiian Shell News 26 (7): 7. Available from https://storage.googleapis.com/conchology-general-images/images/hsn/pdf/1978/7807.pdf [accessed 29 Mar. 2020].

Röckel D. 1982. The continuing uncertainty about Conus siamensis Hwass. Hawaiian Shell News 30 (7): 3. Available from https://storage.googleapis.com/conchology-general-images/images/hsn/pdf/1982/8207.pdf [accessed 29 Mar. 2020].

Röckel D. 1989. Brown cones from the Mediterranean and West Africa. La Conchiglia: International Shell Magazine 238–241: 21–22.

Röckel D., Rolán E. & Monteiro A. 1980a. Cone Shells from Cabo Verde Archipelago a difficult puzzle. Feito, Vigo, Spain.

Röckel D., Cosel R. von & Burnay L.P. 1980b. A new cone from Cape Verde Islands. La Conchiglia: International Shell Magazine 12 (130–131): 10–11.

Röckel D., Korn W. & Kohn A.J. 1995. Manual of the Living Conidae. Vol. I, Indo-Pacific. Christa Hemmen, Wiesbaden. Available from http://theconecollector.com/RKK/RKK_Download.htm [accessed 29 Mar. 2020].

Röding P.F. 1798. Museum Boltenianum, sive, Catalogus Cimeliorum e tribus regnis naturae quae olim collegerat JoA. Fried Bolten, M.D.p. d., per XL. annus Proto physicus Hamburgensis. Pars Secunda continens conchylia sive testacea univalvia, bivalvia and multivalvia. Johan. Christi. Trapii, Hamburg.https://doi.org/10.5962/bhl.title.10588

Rolán E. 1980. Descripción de tres nuevas especies del género Conus Linné, 1758 (Mollusca: Gastropoda), procedentes del archipiélago de Cabo Verde. Bollettino Malacologico 16 (3–4): 79–94.

Rolán E. 1986. Descripción de tres nuevas especies del género Conus (Gastropoda) del archipiélago de Cabo Verde. Publicações Ocasionais da Sociedade Portuguesa de Malacologia 6: 1–16.

Rolán E. 1990. Descripción de nuevas especies y subespecies del género Conus (Mollusca, Neogastro-poda) para el Archipiélago de Cabo Verde. Iberus, Supplement 2: 5–70.

Rolán E. 2005. Malacological Fauna of the Cape Verde Archipelago. ConchBooks, Hackenheim, Germany.

Rolán E. & Raybaudi Massilia G. 2002. Evaluation of character state polarity of Conus radular tooth characters. Bollettino Malacologico Supplemento 4: 175–194.

Rolán E. & Röckel D. 2000. The endemic Conus of Angola. Argonauta 13 (2): 5–44.

Rolán E., Röckel D. & Monteiro A. 1983. New observations on Cape Verde cones, II. La Conchiglia: International Shell Magazine 15 (174–175): 16–17.

Rolán E., Monteiro A. & Fernandes C. 1998. Cone shells from Cape Verde Islands: new developments, with description of a new species. La Conchiglia: International Shell Magazine 30 (286): 36–44.

Rumphius G.E. 1705. D'Amboinsche Rariteitkamer, behelzende eene bescryvinge van allerhande zoo weeke als harde schaalvisschen, te weeten raare Krabben, Kreeften, en diergelyke Zeedieren, als mede allerhande Hoorntjes en Schulpen, die men in d'Amboinsche Zee vindt: daar beneven zommige Mineraalen, Gesteenten, en soorten van Aarde, die in d'Amboinsche, en zommige omleggende Eilanden gevonden worden. Amsterdam. https://doi.org/10.5962/bhl.title.70350

Schröter J.S. 1803. Neue Conchylienarten und Abänderungen, Anmerkungen, und Berichtigungen nach dem linnéischen System der XII. Ausgabe. Archiv für Zoologie und Zootomie 3 (2): 33–78.

Sowerby I G.B. 1823. The Genera of Recent and Fossil Shells, for the use of Students, in Conchology and Geology. London [published between 1821 and 1834]. https://doi.org/10.5962/bhl.title.86281

Sowerby I G.B. 1833. The Conchological Illustrations, or Coloured Figures of all the Hitherto Unfigured Recent Shells. London [published between 1832 and 1841]. https://doi.org/10.5962/bhl.title.11466

Sowerby II G.B. Jr. 1858. Thesaurus Conchyliorum, or Monographs of the Genera of Shells, vol. 3. https://doi.org/10.5962/bhl.title.10596

Tenorio M.J. 2012. Conus ateralbus. The IUCN Red List of Threatened Species 2012: e.T192744A2154590. https://doi.org/10.2305/IUCN.UK.2012-1.RLTS.T192744A2154590.en

Tenorio M.J. & Afonso C.M.L. 2004. Description of four new species of Conus from the Cape Verde Islands (Gastropoda, Conidae.). Visaya 2: 24–37.

Tenorio M.J., Afonso C.M.L. & Rolán E. 2008. New endemic species of Conus (Gastropoda, Conidae) from the Islands of São Nicolau, Santo Antão and Sal in the Cape Verde Archipelago. Vita Malacologica 6: 1–10.

Tenorio M.J., Afonso C.M.L., Cunha R.L. & Rolán E. 2014. New species of Africonus (Gastropoda, Conidae) from Boa Vista in the Cape Verde Archipelago: molecular and morphological characterization. Xenophora Taxonomy 2: 5–18.

Tenorio M.J., Lorenz F. & Dominguez M. 2016. New insights into Conus jourdani da Motta, 1984 (Gastropoda, Conidae), an endemic species from Saint Helena Island. Xenophora Taxonomy 11: 32–42.

Tenorio M.J., Afonso C.M.L., Rolán E., Pires S., Vasconcelos P., Abalde S. & Zardoya R. 2017. DNA sequences disclose a new species of Africonus cone snail from São Vicente (Gastropoda: Conidae). Zoologia Caboverdiana 6 (2): 32–41. Available from http://www.scvz.org/zoolcv/vol6no2/ZoolCV2017_6_2_Tenorio_etal_Africonus%20freitasi.pdf [accessed 10 May 2020].

Tenorio M.J., Abalde S. & Zardoya R. 2018. Identification of new species of Kalloconus and Africonus (Gastropoda, Conidae) from the Cabo Verde Archipelago through mitochondrial genome comparison. The Festivus 50 (2): 73–88.

Trovão H.F.M. 1978. Contribuição para o estudo dos moluscos Gasterópodes da familia Conidae de Angola. Novas espécies de Conus Linné, 1758 de Angola (Mollusca: Gastropoda). Boletim do Centro Português de Actividades Subaquáticas ser. 4, 4: 11–20.

Trovão H.F.M. 1979. Contribuição para o estudo dos Moluscos Gasterópodos da famaia Conidae da África occidental. Amphitrite 1 (1): 3–12.

Trovão H.F.M. & Rolán E. 1986. Description of a new species for the genus Conus (Gastropoda: Conidae), from the Cape Verde Islands. Publicações Ocasionais da Sociedade Portuguesa de Malacologia 7: 9–15.

Trovão H.F.M., Rolán E. & Félix-Alves A.V. 1990. A new species of Conus from the Cabo Verde Archipelago, West Africa (Mollusca: Gastropoda: Conidae). Publicações Ocasionais da Sociedade Portuguesa de Malacologia 15: 69–78.

Tucker J.K. & Tenorio M.J. 2009. Systematic Classification of Recent and Fossil Conoidean Gastropods. ConchBooks, Hackenheim, Germany.

Tucker J.K. & Tenorio M.J. 2013. Illustrated Catalog of the Living Cone Shells. MdM Publishing, Wellington, Florida, USA.

Uribe J.E, Puillandre N. & Zardoya R. 2017. Beyond Conus: phylogenetic relationships of Conidae based on complete mitochondrial genomes. Molecular Phylogenetics and Evolution 107: 142–151. https://doi.org/10.1016/j.ympev.2016.10.008

van Rossum H.M. 1996. A new Conus species from the West African Coast. World Shells 12: 59–61.

van Rossum H.M. 1997. The Conus pulcher Lightfoot, 1786 complex with description of two new forms. World Shells 20: 36–42.

Wagner R.J.L. & Abbott R.T. 1978. Wagner and Abbott’s Standard Catalog of Shells. Third Edition with Supplements. American Malacologists Inc., Conidae section, Greenville, Delaware, USA.

Walls J.G. 1978. Diagnoses of four new cones (Mollusca: Conidae). The Pariah 2: 1–7.

Weinkauff H.C. 1873. Die familie der Coneae oder Conidae. Systematisches Conchylien-Cabinet von Martini und Chemnitz vol. 4 (2): 125–244. Bauer & Raspe, Nürnberg.

Wilson, J. 1831. Illustrations of Zoology, being Representations of New, Rare or Remarkable Subjects of the Animal Kingdom...with...Descriptive Details. Edinburgh [published between 1826 and 1831].

WoRMS Editorial Board. 2019. World register of marine species. Available from http://www.marinespecies.org [accessed March 2019].

Published
2020-06-10
How to Cite
Tenorio, M. J., Abalde, S., Pardos-Blas, J. R., & Zardoya, R. (2020). Taxonomic revision of West African cone snails (Gastropoda: Conidae) based upon mitogenomic studies: implications for conservation. European Journal of Taxonomy, (663). https://doi.org/10.5852/ejt.2020.663