Abstract
So far, of the 292 known species of solenogasters (Mollusca, Aplacophora), 62 belong to the clade Pholidoskepia Salvini-Plawen, 1978. Of these, only two have an abyssal distribution (3500–6000 m depth). Among Pholidoskepia, Dondersiidae Simroth, 1893 is the most diverse family. This study contributes to the knowledge of this family with the description of one new genus and six new species from the abyssal South Atlantic Ocean: Dondersia ? foraminosa sp. n., Nematomenia divae sp. n., Nematomenia brasiliensis sp. n., Nematomenia ? guineana sp. n., Helluoherpia vieiralaneroi sp. n. and Inopinatamenia (gen. n.) calamitosa sp. n. Specimens were collected during DIVA (Latitudinal Gradients of Deep-Sea BioDIVersity in the Atlantic Ocean) expeditions in the Guinea (DIVA 2 Me 63/2, 2005) and Brazil (DIVA 3 Me 79/1, 2008) Basins. Specimens were characterized based primarily on the sclerites and internal anatomy, which was studied using histology. The importance of the radula and mantle sclerites for taxonomy is emphasized. Amended diagnoses for the family and some genera within this family are provided. This contribution increases the described diversity of Dondersiidae to ten genera and 38 species and highlights the need for more study of solenogasters in the deep sea.
References
Bergmeier, F.S., Haszprunar, G., Todt, C. & Jörger, K.M. (2016) Lost in a taxonomic Bermuda Triangle: comparative 3D-microanatomy of cryptic mesopsammic Solenogastres (Mollusca). Organisms Diversity & Evolution,16 (3), 613–639.
https://doi.org/10.1007/s13127-016-0266-6
Bergmeier, F.S., Brandt, A., Schwabe, E. & Jörger, K.M. (2017) Abyssal Solenogastres (Mollusca, Aplacophora) from the Northwest pacific: scratching the surface of Deep-Sea diversity using integrative taxonomy. Frontiers in Marine Science, 4, 410.
https://doi.org/10.3389/fmars.2017.00410
Bergmeier, F.S., Haszprunar, G., Brandt, A., Saito, H., Kano, Y. & Jörger, K. (2019) Of basins, plains and trenches: systematics and distribution of Solenogastres (Mollusca, Aplacophora) in the Northwest Pacific. Progress in Oceanography, 178, 102–187.
https://doi.org/10.1016/j.pocean.2019.102187
Büchinger, T. & Handl, C. (1996) Squamatoherpia tricuspidata gen. n. et sp. n. aus der Nordsee (Mollusca: Solenogastres: Dondersiidae). Annalen des Naturhistorischen Museums in Wien, 98B, 57–63.
Cobo, M.C. & Kocot, K.M. (2020) Micromenia amphiatlantica sp. nov.: first solenogaster (Mollusca, Aplacophora) with an amphi-Atlantic distribution and insight into abyssal solenogaster diversity. Deep-Sea Research, Part I, 157, 103–189.
https://doi.org/10.1016/j.dsr.2019.103189
Cobo, M.C., Pedrouzo, L., García-Álvarez, O., Urgorri, V., Barrio, L. & Cobo, F. (2013) Contribution to abyssal Angola Basin Solenogastres (Mollusca) with preliminary records of two species of the order Pholidoskespia. Açoreana, Supplement 8, 162.
Folmer, O., Hoeh, W. R., Black, M.B. & Vrijenhoek, R.C. (1994) Conserved primers for PCR amplification of mitochondrial DNA from different invertebrate phyla. Molecular Marine Biology and Biotechnology, 3 (5), 294–299.
García-Álvarez, O. & Salvini-Plawen, L. (2007) Species and diagnosis of the families and genera of Solenogastres (Mollusca). Iberus, 25 (2), 73–143.
García-Álvarez, O., Salvini-Plawen, L. & Urgorri, V. (2014) Solenogastres. In: García-Álvarez, O., Salvini-Plawen, L., Urgorri, V., Souza Troncoso, J. (Eds.), Mollusca Solenogastres, Caudofoveata, Monoplacophora. Fauna Iberica. Vol. 38. Museo Nacional de Ciencias Naturales. CSIC, Madrid, pp. 11–26.
García-Álvarez, O., Urgorri, V. & Cristobo, F.J. (2000) Synopsis of the interstitial Solenogastres (Mollusca). Argonauta, 14 (2), 27–37.
Gil-Mansilla, E., García-Álvarez, O. & Urgorri, V. (2008) Metodología para la recolección, conservación y estudio de los moluscos Solenogastros. Sociedad Española de Malacología. Reseñas Malacológicas, XIII, 1–31.
Gil-Mansilla, E., García-Álvarez, O. & Urgorri, V. (2009) A new genus and two new species of Simrothiellidae (Solenogastres: Cavibelonia) from the Abyssal Angola Basin. Journal of the Marine Biological Association of the United Kingdom, 89 (7), 1507–1515.
https://doi.org/10.1017/S0025315409000666
Gil-Mansilla, E., García-Álvarez, O. & Urgorri, V. (2011) A new genus and two new species of Cavibelonia (Mollusca: Solenogastres) from the Abyssal Angola Basin. Cahiers de Biologie Marine, 52, 233–244.
Gil-Mansilla, E., García-Álvarez, O. & Urgorri, V. (2012) Three new species of Kruppomenia (Solenogastres: Simrothiellidae) from the abyssal Angola Basin. Journal of the Marine Biological Association of the United Kingdom, 92 (2), 349–335.
https://doi.org/10.1017/S0025315411001275
Handl, C.H. & Büchinger, T. (1996) Helluoherpia aegiri gen. n. et sp. n. aus Norwegen (Mollusca: Solenogastres: Dondersiidae). Annalen des Naturhistorischen Hofmuseums in Wien, 98, 65–70.
Handl, C.H., Salvini-Plawen, L. & Høisæter, T. (2001) New records of Solenogastres-Pholidoskepia (Mollusca) from Norwegian fjords and shelf waters including two new species. Sarsia, 86 (4–5), 367–381.
https://doi.org/10.1080/00364827.2001.10425524
Handl, C.H. & Todt, C. (2005) Foregut glands of Solenogastres (Mollusca): anatomy and revised terminology. Journal of Morphology, 265, 28–42.
https://doi.org/10.1002/jmor.10336
Heath, H. (1911) Reports on the scientific results of the expedition to the tropical Pacific, XIV. The Solenogastres. Memoirs of the Museum of Comparative Zoology at Harvard College, 45 (1), 1–179.
Heath, H. (1918) Solenogastres from the eastern coast of North America. Memoirs of the Museum of Comparative Zoology at Harvard College, 45 (2), 183–263.
https://doi.org/10.5962/bhl.title.49189
Hubrecht, A.W. (1888) Dondersia festiva gen. et spec. nov. In: Feestbundel aan Franciscus Cornelis Donders op den 27sten. Mei 1888, aangeboden door het Nederlandsch Tijdschrift voor Geneeskunde. F. van Rossen, Amsterdam, pp. 324–339.
Klink, S.P., Bergmeier, F.S., Neusser, T.P. & Jörger, K.M. (2015) Stranded on a lonely island: Description of Dondersia (?) todtae sp. nov., the first shelf solenogaster (Mollusca, Aplacophora) from the Azores. Açoreana, 10 (4), 603–618.
Kocot, K.M., Cannon, J.T., Todt, C., Citarella, M.R., Kohn, A., Meyer, A., Santos, S.R., Schander, C., Moroz, L.L., Lieb, B. & Halaynch, K.M. (2011) Phylogenomics reveals deep molluscan relationships. Nature, 477, 452–456.
https://doi.org/10.1038/nature10382
Kocot, K.M. & Todt, C. (2014) Three new meiofaunal solenogaster species (Mollusca: Aplacophora) from the north-east Pacific. Journal of Natural History, 48 (45–48), 3007–3031.
https://doi.org/10.1080/00222933.2014.961987
Kocot, K.M., Todt, C., Mikkelsen, N. & Halanych, K. (2019) Phylogenomics of Aplacophora (Mollusca, Aculifera) and a solenogaster without a foot. Proceedings of the Royal Society B: Biological Sciences, 286 (1902), 20190115.
https://doi.org/10.1098/rspb.2019.0115
Kocot, K.M., Poustka, A.J, Stöger, I., Halanych, K.M. & Schrödl, M. (2020) New data from Monoplacophora and a carefully-curated dataset resolve molluscan relationships. Scientific Reports, 10 (1), 1–8.
https://doi.org/10.1038/s41598-019-56728-w
Kowalevsky, A. (1881) Neomenia coralliophila i Ceoloplana metschnikowii. Izvestia Obshchestvo Liubiteley. Estestvozaniya Anthropologii i Etnografii (Comunicazioni della Società Imperiale degli Amici delle Scienze Naturali), Moskwa, 43 (1), 1–5.
Leloup, E. (1948) Un nouveau Solénogastre a peau nue de l’Océan Glaciar Arctique, Micromenia simplex g. nov., sp. nov. Bulletin du Muséum d’Histoire Naturelle de Belgique, 24 (19), 1–7.
Marion, A. & Kowalevsky, A. (1886) Organisation du Lepidomenia hystrix, nouveau type de Solénogastre. Comptes Rendus de l’Académie des Sciences Paris, 103 (17), 757–759.
Morse, M.P. & Scheltema, A.H. (1988) Aplacophora. In: Higgins, R.P. & Thiel, H. (Eds.), Introduction to the study of Meiofauna. Smithsonian Press, Washington, pp. 447–450.
Nierstrasz, H. (1902) The Solenogastres of the Siboga-Expedition. Siboga-Expeditie, 47, 1–46.
https://doi.org/10.5962/bhl.title.10915
Nierstrasz, H. (1908) The Solenogastres of Discovery-Expedition. National Antarctic Expeditions 1901–1904. Natural History, 4, 38–46.
Odhner, N.H. (1920) Norwegian Solenogastres. Bergens Museum Aarbok (1918–1919), 1 (3), 1–86.
Ostermair, L., Brandt, A., Haszprunar, G., Jörger, K.M. & Bergmeier, F.S. (2017) First insights into the solenogaster diversity of the Sea of Okhotsk with the description of a new species of Kruppomenia (Simrothiellidae, Cavibelonia). Deep sea research Part II: topical studies in oceanography, 154, 214–229.
https://doi.org/10.1016/j.dsr2.2017.12.008
Pilsbry, H.A. (1897–1898). Manual of conchology, structural and systematic, with illustrations of the species. In: Scaphopoda, by H.A. Pilsbry & B. Sharp. Aplacophora. Index to genera and subgenera. Vols. II—XVII. Conchological section, Academy of Natural Sciences of Philadelphia, Philadelphia, pp. 1–32.
Pruvot, G. (1890) Sur quelques Néoméniées nouvelles de la Méditerranée. Archives de Zoologie Expérimentale et générale, 2 (8), 21–24.
Pruvot, G. (1891) Sur l’organisation de quelques néomeniens des côtes de France. Archives de Zoologie Expérimentale et générale, Série 2, 9, 699–810.
Pruvot, G. (1899) Sur deux Néoméniens nouveaux de la Méditerranée. Archives de Zoologie Expérimentale et Générale, 3 (7), 461–509.
Pruvot, G. (1902) Sur les affinités et le classement des néoéniens. Archives de Zoologie Expérimentale et Générale, 3, 10.
Salvini-Plawen, L. (1972) Revision der monegassischen Solenogastres (Mollusca, Aculifera). Zeitschrift für Zoologische Systematik und Evolutionsforschung, 10 (3), 215–240.
https://doi.org/10.1111/j.1439-0469.1972.tb00799.x
Salvini-Plawen, L. (1978) Antarktische und Subantarktische Solenogastres - Eine monographie: 1898–1974. Zoologica, 128, 1–315.
Salvini-Plawen, L. (1985a) Early evolution and the Primitive Groups. In: Trueman, E.R. & Clarke, M.R. (Eds.), The Mollusca. Vol. 10. Evolution. Academy Press. Orlando, pp. 59–150.
https://doi.org/10.1016/B978-0-12-751410-9.50010-1
Salvini-Plawen, L. (1985b) New interstitial Solenogastres (Mollusca). Stygologia, 1 (1), 101–108.
Salvini-Plawen, L. (1988) Einige Solenogastres (Mollusca) der europäische Meiofauna. Annalen des Naturhistorischen Museum in Wien, 90B, 373–385.
Salvini-Plawen, L. (2003a) Contributions to West-Mediterranean Solenogastres (Mollusca) with three new species. Iberus, 21 (2), 37–60.
Salvini-Plawen, L. (2003b) On the phylogenetic significance of the aplacophoran Mollusca. Iberus, 21 (1), 67–97.
Salvini-Plawen, L. (2008) Three new species of Simrothiellidae (Solenogastres) associated with the hot-vent biotope. Journal of Molluscan Studies, 74 (3), 223–238.
https://doi.org/10.1093/mollus/eyn010
Scheltema, A.H. (1988) Class Aplacophora. In: Scott, P.V. & Blake, J.A. (Eds.), Taxonomic of the Benthic Fauna of the Santa Maria Basin and Western Santa Barabara Channel. Vol 8. The mollusca Part 1. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 3–47.
Scheltema, A.H. (1999) New eastern Atlantic neomenioid aplacophoran molluscs (Neomeniamorpha, Aplacophora). Ophelia, 5, 1–28.
https://doi.org/10.1080/00785326.1999.10409397
Scheltema, A.H. & Schander, C. (2000) Discrimination and phylogeny of solenogaster species through the morphology of hard parts (Mollusca, Aplacophora, Neomeniomorpha). Biological Bulletin, 198, 121–151.
https://doi.org/10.2307/1542810
Scheltema, A.H., Schander, C. & Kocot, K.M. (2012) Hard and soft anatomy in two genera of Dondersiidae (Mollusca, Aplacophora, Solenogastres). Biological Bulletin, 222 (3), 233–269.
https://doi.org/10.1086/BBLv222n3p233
Schwabl, M. (1955) Rupertomenia fodiens nov. gen., nov. spec., eine neue Lepidomeniide von der Südwestküste Schwedens. Österreichische Zoologische Zeitschrift, 6 (1–2), 90–146.
Simroth, H. (1893) Aplacophora. In: Bronn, H.G. (Ed.), Die Klassen und Ordnungen des Thier-Reichs, 3 (1), 128–233.
Smith, S., Wilson, N.G., Goetz, F., Feehery, C., Andrade, S.C.S., Rouse, G.W., Giribet, G. & Dunn, C.W. (2011) Resolving the evolutionary relationships of molluscs with phylogenomic tools. Nature, 480, 364–367.
https://doi.org/10.1038/nature10526
Thiele, J. (1913a) Antarktische Solenogastren. Deutsche Südpolar-Expedition, 1901–1903, im Auftrage des Reichsamtes des Innern, 14. Zoologie, 6 (1), 35–65.
Thiele, J. (1913b) Ein neuer Solenogaster von Spitzbergen. Sitzungsberichte der Gesellschaft Naturforschender Freunde Berlin, 1913 (2), 160–162.
Thiele, J. (1913c) Mollusca: Solenogastres. In: Schulze, F.E. (Ed.), Das Tierreich. Eine Zusammenstellung und Kennzeichnung der rezenten Tierformen. 38 R. Friedlander & Sohn, Berlin, 57 pp.
Todt, C. (2013) Aplacophoran mollusks—still obscure and difficult? American Malacological Bulletin, 31 (1), 181–187.
https://doi.org/10.4003/006.031.0110