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Type: Article
Published: 2020-07-15
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Reinstatement of species belonging Marphysa sanguinea complex (Annelida: Eunicidae) and description of new species from the mid-Pacific Ocean and the Adriatic Sea

Estructura y Función del Bentos, Depto. Sistemática y Ecología Acuática, El Colegio de la Frontera Sur, Chetumal, Quintana Roo, México. South China Sea Repository and Reference Centre, Institute of Oceanography and Environment, Universiti Malaysia Terengganu, 21030 Kuala Nerus, Terengganu, Malaysia.
South China Sea Repository and Reference Centre, Institute of Oceanography and Environment, Universiti Malaysia Terengganu, 21030 Kuala Nerus, Terengganu, Malaysia.
Annelida Branchiae pectinate chaetae parapodia lobes redescription synonymy

Abstract

Marphysa sanguinea Montagu, 1813, the type species of genus Marphysa, was described with few characteristics and subsequently reported in tropical and temperate waters as a cosmopolitan species. Recent comparative studies have shown that M. sanguinea has a limited distribution to the Northeast Atlantic. As a result, species from the tropical and subtropical zones are now being redescribed as new species. However, this issue is not completely resolved because of the taxonomic status of seven nominal species, considered as junior synonymies of M. sanguinea. Herein, we examined the type and additional materials from five species in synonymy M. sanguinea and performed a compared analysis with the neotype of M. sanguinea. Additionally, we reviewed specimens from two localities in the mid-Pacific Ocean and the Adriatic Sea, where M. sanguinea was previously reported. As a result of this study, three species were reinstated (M. californica Moore, 1909, M. leidii de Quatrefages, 1866, and M. parishii Baird, 1869) and two subspecies, are now raised to species level (M. americana n. status and M. brevibranchiata n. status). In addition, two new species from Hawaii and Croatia are described: M. baileybrockae n. sp. and M. birgeri n. sp., respectively. Morphological comparisons with most species currently assigned to the M. sanguinea complex are presented. This work provides more support for rejecting the hypothesis that M. sanguinea is a cosmopolitan species and we encourage the use of novel and traditional morphological characters to differentiate species within the complex.

 

References

  1. Abbott, D.P. (1946) Some polychaetous annelids from a Hawaiian fish pond. University of Hawaii research publications, 23, 1–24.

    Abe, H., Tanaka, M., Taru, M., Abe, S. & Nishigaki, A. (2019) Molecular evidence for the existence of five cryptic species within the Japanese species of Marphysa (Annelida: Eunicidae) known as “Iwa-mushi. Plankton and Benthos Research, 14 (4), 303–314.

    https://doi.org/10.3800/pbr.14.303

    Andrews, E.A. (1891) Report upon the Annelida Polychaeta of Beaufort, North Carolina. Proceedings of the United States National Museum,14 (852), 277–302.

    https://doi.org/10.5479/si.00963801.14-852.277

    Audouin, J.V. & Milner-Edwards, H. (1834) Recherches pour servir a l’histoire naturelle du littoral de la France, ou, Recueil de mémoires sur l’anatomie, la physiologie, la classification et les moeurs des animaux des nos côtes: ouvrage accompagn de planches faites d’après nature. Tome 2. Annélides. Part 1. Crochard, libraire, Paris, 290 pp.

    Augener, H. (1931) Die bodensässigen Polychäten nebst einer Hirudinee der Meteor-Fahrt. Mitteilungen der Zooologisches Staatinstitut und zoologisches Museum, Hamburg, 44, 279–313.

    Baird, W. (1869) Remarks on several genera of annelides, belonging to the group Eunicea, with a notice of such species as are

    contained in the collection of the British Museum, and a description of some others hitherto undescribed. Journal of the Linnean Society of London, 10, 341–361.

    https://doi.org/10.1111/j.1096-3642.1869.tb00665.x

    Bellan, G. (1964) Contribution l’étude systématique et écologique des annélides polychètes de la Méditerranée. Recueil des Travaux de la Station Marine d’Endoume, 49, 1–371.

    Berthold, A.A. (1827) Latreille’s Natürliche Familien des Thierreichs: Aus dem Franzosischen, mit Anmerkungen und Zusätzen. Verlage Landes-Industrie-Comptoirs, Weimar, 606 pp.

    Carrera-Parra, L.F. (2009) Eunicidae Berthold, 1827. In: de León-González, J.A., Bastida-Zavala, J.R., Carrera-Parra, L.F., García-Garza, M.E., Peña-Rivera, A., Salazar-Vallejo, S.I. & Solís-Weiss, V. (Eds.), Poliquetos (Annelida: Polychaeta) de México y América Tropical. Universidad Autónoma de Nuevo León, Monterrey, pp. 165–181.

    Carrera-Parra, L.F. & Salazar-Vallejo, S.I. (2011) Redescriptions of Eunice filamentosa and E. denticulata and description of E. tovarae n. sp. (Polychaeta: Eunicidae), highlighted with morphological and molecular data. Zootaxa, 2880 (1), 51–64.

    https://doi.org/10.11646/zootaxa.2880.1.5

    Cuvier, G. (1817) Le régne animal distribu d’après son organisation, pour servir de base l’histoire naturelle des animaux et d’introduction l’anatomie comparée: Les Reptiles, les Poisson, les Mollusques et les Annélides. Deterville Libraire, Paris, 532 pp.

    Dales, R.P. (1962) The polychaete stomodeum and the inter-relationships of the families of Polychaeta. Proceedings of the Zoological Society of London, 139, 389–428.

    https://doi.org/10.1111/j.1469-7998.1962.tb01837.x

    Day, J.H. (1967) A Monograph on the Polychaeta of Southern Africa. Part I. Errantia. Bulletin of the British Museum (Natural History), 1967, 1–458.

    https://doi.org/10.5962/bhl.title.8596

    de Blainville, H.M.D. de. (1825) NERÉIDE Nereis (Chétopodes ), [alphabetical dictionary entry]. In: [multi-volume book] Dictionnaire des Sciences naturelles, dans lequel on traite méthodiquement des différens êtres de la nature, considérés soit en eux-mêmes, d’apres l’état actuel de nos connaisance, soit relativement a l’utilit qu’en peuvent retirer la médicine, l’agriculture, le commerce et les arts. Suivi d’une biographie des plus célèbres naturalistes, 34, pp. 408–455.

    de Blainville, H. (1828a) Vers. In: Dictionnaire des Sciences naturelles, dans lequel on traite methodiquement des differens etres de la nature, consideres soit en eux-memes. d’apres l’etat actuel de nos connais sciences, soit relativement a l’utilite qu’en peuvent retirer la medicine, l’agriculture, le commerce et les arts. Suivie d’une biographie des plus celebres naturalistes, 57, pp. 365–625.

    de Blainville, H.M.D. (1828b) Mollusques, Vers et Zoophytes. In: Dictionnaire des Sciences naturelles, dans lequel on traite methodiquement des differens etres de la nature, consideres soit en eux-memes. d’apres l’etat actuel de nos connais sciences, soit relativement a l’utilite qu’en peuvent retirer la medicine, l’agriculture, le commerce et les arts. Suive d’une biographie des plus celebres naturalistes, 57 (Tome LVII. Vea–Vers), pp. 365.

    de Quatrefages, A. (1865a) Note sur la classification des Annélides. Compte rendu hebdomadaire des séances de l’Academie des sciences, 60, 586–600.

    de Quatrefages, A. (1865b) Note sur la Classification des Annelides. Annales des Sciences Naturelles, Paris, Zoologie, 3, 253–296.

    de Quatrefages, A. (1866 [1865]) Histoire Naturelle des Annelés Marins et d’Eau Douce: Annélides et Géphyriens. Tome Premier. Collection des Suites a Buffon formant avec les Oeuvres de cet auteur un Cours Complet d’Histoire Naturelle. Librairie Encyclopédique de Roret, Paris 588 pp. [Wright, E.P. (1866) Annelida. Zoological Record, Section 6 (Annelida, etc.), 578–600 pp.]

    https://doi.org/10.5962/bhl.title.122818

    Elgetany, A.H., El-Ghobashy, A.E., Ghoneim, A.M. & Struck, T.H. (2018) Description of a new species of the genus Marphysa (Eunicidae), Marphysa aegypti n. sp., based on molecular and morphological evidence. Invertebrate Zoology, 15 (1), 71–84.

    https://doi.org/10.15298/invertzool.15.1.05

    Fauchald, K. (1970) Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitimidae, Arabellidae, Lysaretidae and Dorvilleidae from Western Mexico. Allan Hancock Monographs in Marine Biology, 5, 1–335.

    Fauchald, K. (1977) The polychaete worms, definitions and keys to the orders, families and genera. Natural History Museum of Los Angeles Countya 28, 1–188.

    Fauchald, K. (1991) A morphometric study of eunicids polychaetes from Belize, Western Caribbean Sea. Ophelia, Supplement 5, 47–53.

    Fauvel, P. (1911) Troisième note préliminaire sur les polychètes provenant des campagnes de l’Hirondelle et de la Princesse-Alice, ou déposées dans la Musée Océanographique de Monaco. Bulletin de l’Institut Océanographique de Monaco, 194, 1–41.

    Fauvel, P. (1923) Polychètes errantes. Faune de France. Librairie de la Faculte des Sciences, Paris, 5, 1–488.

    Fauvel, P. (1936) Annelides Polychetes du Japon. Mem. Memoirs of the College of Science, Kyoto Imperial University, 12, 41–92.

    Glasby, C.J. & Hutchings, P.A. (2010) A new species of Marphysa Quatrefages, 1865 (Polychaeta: Eunicida: Eunicidae) from northern Australia and a review of similar taxa from the Indo-west Pacific, including the genus Nauphanta Kinberg, 1865. Zootaxa, 2352 (1), 29–45.

    https://doi.org/10.11646/zootaxa.2352.1.2

    Glasby, C.J., Mandario, M.A., Burghardt, I., Kupriyanova, E., Gunton, L.M. & Hutchings, P.A. 2019. A new species of the Marphysa sanguinea Quatrefages, 1865 species complex (Annelida: Eunicida: Eunicidae) from the Philippines, and a key to members of the complex in Southeast Asia. Zootaxa, 4674 (2), 264–282.

    https://doi.org/10.11646/zootaxa.4674.2.7

    Grube, A.E. (1864) Die insel Lussin und ihre Meeresfauna: Nach einem Sechswöchentlichen Aufenthalte. Ferdinand Hirt, Breslau, 116 pp.

    Grube, A.E. (1878) Untersuchungen ueber die Familie Eunicea. Jahres-Bericht der Schlesischen Gesellschaft für Vaterländische Cultur, Berlin, 1878, 37–62.

    Grube, A.E. (1881) Beschreibungen von neuen Anneliden des zoologischen Museums zu Berlin. Sitzungsberichte der Gesellschaft der naturforschende Freunde zur Berlin, 19 (7), 109–117.

    Hansen, A. (1882) Recherches sur les annélides recueillies par M. le professeur Édouard van Benedon pendant son voyage au Brésil et la Plata. Mémoires Couronnes et Mémoires des Savants Etrangers publiés par L’Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique, 44 (3), 1–29.

    Hartman, O. (1944a) Polychaetous Annelids. Part V. Eunicea. Allan Hancock Pacific Expedition, 10 (1), 1–237.

    Hartman, O. (1944b) New England Annelida. Pt. 2, including the unpublished plates by Verrill with reconstructed captions. Bulletin of American Museum of Natural History, 82 (7), 327–344.

    Hartman, O. (1945) The marine annelids of North Carolina. Bulletin of Duke University Marine Station, 2, 1–51.

    Hartman, O. (1956) Polychaetous annelids erected by Treadwell, 1891 to 1948, together with a brief chronology. Bulletin of American Museum of Natural History, 109 (2), 239–310.

    Hartman, O. (1961) Polychaetous annelids from California. Allan Hancock Pacific Expedition, 25, 1–226.

    Hartman, O. (1966) Polychaetous annelids of the Hawaiian Islands. Occasional Papers of the Bernice P. Bishop Museum, 23 (11), 163–252.

    Hartmann-Schröder, G. (1965) Die Polychaeten des Sublitorals. In: Hartmann-Schröder G, Hartmann G (Eds) Zur Kenntnis des Sublitorals der chilenischen Küste unter besonderer Berücksichtigung der Polychaeten und Ostracoden. (Mit bemerkungen über den Einfluss sauerstoffarmer Strömungen auf die Besiedlung von marien Sedimenten.). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 62, pp. 59–305.

    Hutchings, P.A. & Karageorgopoulos, P. (2003) Designation of a neotype of Marphysa sanguinea (Montagu, 1813) and a description of a new species of Marphysa from eastern Australia. Hydrobiologia, 496, 87–94.

    https://doi.org/10.1023/A:1026124310552

    Hutchings, P., Glasby, C.J. & Wijnhoven, S. (2012) Note on additional diagnostic characters for Marphysa sanguinea (Montagu, 1813) (Annelida: Eunicida: Eunicidae), a recently introduced species in the Netherlands. Aquatic Invasions, 7 (2), 277–282.

    https://doi.org/10.3391/ai.2012.7.2.014

    International Commission on Zoological Nomenclature [ICZN] (1999) International code of zoological nomenclature. 4th Edition. The International Trust for Zoological Nomenclature, London, 306 pp.

    Izuka, A. (1907) On two new species of annelids belonging to the Euncidae. Zoological Magazine (Dobutsugasku zasshi), 19, 139–143. [translated into English]

    Kara, J., Molina-Acevedo, I.C., Zanol, J., Simon, C.A. & Idris, I. (2020) Morphological and molecular systematic review of Marphysa Quatrefages, 1865 (Annelida: Eunicidae) species from South Africa. PeerJ. [in review]

    Lamarck, J.B.P.A. (1818) Histoire naturelle des Animaux sans Vertèbres, préséntant les caractères généraux et particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres, et la citation des principales espèces qui s’y rapportent; précédée d’une Introduction offrant la Détermination des caractères essentiels de l`Animal, sa distinction du végétal et des autres corps naturels, enfin, l’Exposition des Principes fondamentaux de la Zoologie. Déterville and Verdière Libraire, Paris, 612 pp.

    Lavesque, N., Daffe, G., Bonifácio, P. & Hutchings, P. (2017) A new species of the Marphysa sanguinea complex from French waters (Bay of Biscay, NE Atlantic) (Annelida, Eunicidae). ZooKeys, 716, 1–17.

    https://doi.org/10.3897/zookeys.716.14070

    Lavesque, N., Daffe, G., Grall, J., Zanol, J., Gouillieux, B. & Hutchings, P. (2019) Guess who? On the importance of using appropriate name: case study of Marphysa sanguinea (Montagu, 1813). ZooKeys, 859, 1–15.

    https://doi.org/10.3897/zookeys.859.34117

    Lewis, C. & Karageorgopoulos, P. (2008) A new species of Marphysa (Eunicidae) from the western Cape of South Africa. Journal of marine biological Association United Kingdom, 88, 277–287.

    https://doi.org/10.1017/S002531540800009X

    Leidy, J. (1855) Contributions towards a knowledge of the marine invertebrate fauna of the coasts of Rhode Island and New Jersey. Journal of the Academy of Natural Sciences of philadelphia, New Series, 3, 135–152.

    Liu, Y., Hutchings, P.A. & Sun, S. (2017) Three new species of Marphysa Quatrefages, 1865 (Polychaeta: Eunicida: Eunicidae) from the south coast of China and redescription of Marphysa sinensis Monro, 1934. Zootaxa, 4263 (2), 228–250.

    https://doi.org/10.11646/zootaxa.4377.2.3

    Liu, Y., Hutchings, P.A. & Kupriyanova, E. (2018) Two new species of Marphysa Quatrefages, 1865 (Polychaeta: Eunicida: Eunicidae) from northern coast of China and redescription for Marphysa orientalis Treadwell, 1936. Zootaxa, 4377 (2), 191–215.

    https://doi.org/10.11646/zootaxa.4377.2.3

    Malmgren, A.J. (1867) Annulata Polychaeta Spetsbergi, Grœnlandi, Islandi et Scandinavi. Hactenus Cognita. Ex Officina Frenckelliana, Helsingforsl, 127 pp.

    https://doi.org/10.5962/bhl.title.13358

    Martin, D., Gil, J., Zanol, J., Meca, M.A. & Pérez Portela, R. (2020) Digging the diversity of Iberian bait worms Marphysa (Annelida, Eunicidae). PLoS ONE, 15 (1), e0226749.

    https://doi.org/10.1371/journal.pone.0226749

    McIntosh, W.C. (1910) A monograph of the British annelids. Polychaeta. Syllidae to Ariciidae. Ray Society of London, 2 (2), 233–524.

    Miura, T. (1986) Japanese polychaetes of the genera Eunice and Euniphysa: Taxonomy and branchial distribution patterns. Publications of the Seto Marine Biological Laboratory, 31 (3), 269–325.

    https://doi.org/10.5134/176125

    Miura, T. (1977) Eunicid polychaetous annelids from Japan, II. La Mer, 15, 11–31.

    Molina-Acevedo, I.C. & Carrera-Parra, L.F. (2015) Reinstatement of three species of the Marphysa sanguinea complex (Polychaeta: Eunicidae) from the Grand Caribbean Region. Zootaxa, 3925 (1), 37–55.

    https://doi.org/10.11646/zootaxa.3925.1.3

    Molina-Acevedo, I.C. & Carrera-Parra, L.F. (2017) Revision of Marphysa de Quatrefages, 1865 and some species of Nicidion Kinberg, 1865 with the erection of a new genus (Polychaeta: Eunicidae) from the Grand Caribbean. Zootaxa, 4241 (1), 1–62.

    https://doi.org/10.11646/zootaxa.3925.1.3

    Molina-Acevedo, I.C. (2018) Morphological revision of the Subgroup 1 Fauchald, 1970 of Marphysa de Quatrefages, 1865 (Eunicidae: Polychaeta). Zootaxa, 4480 (1), 001–125.

    https://doi.org/10.11646/zootaxa.4480.1.1

    Molina-Acevedo, I.C. (2019) Redescription of two species previously regarded as Marphysa de Quatrefages, 1865, with the description of new species of Treadwellphysa Molina-Acevedo and Carrera-Parra, 2017. Journal of Natural History, 53 (9–10), 517–540.

    https://doi.org/10.1080/00222933.2019.1596328

    Monro, C.C.A. (1933) The Polychaeta Errantia collected by Dr. C. Crossland at Colón, in the Panama Region, and the Galapagos Islands during the expedition of the S.Y. ‘St. George’. Proceedings of the Zoological Society of London, 1933, 1–196.

    https://doi.org/10.1111/j.1096-3642.1933.tb01578.x

    Montagu, G. (1813) Descriptions of several new or rare animals principally marine, found on the south coast of Devonshire. Transactions of the Linnean Society of London, 11, 18–21.

    https://doi.org/10.1111/j.1096-3642.1813.tb00035.x

    Moore, J.P. (1909) Polychaetous annelids from Monterey Bay and San Diego, California. Proceedings of the Academy of Natural Sciences of Philadelphia, 61, 235–295.

    Nishi, E. & Tanaka, K. (2011) Phylum Annelida (Classes Polychaeta and Myzostomida): a catalogue of Akira Izuka’s type and non-type polychaete collection in the University Museum, the University of Tokyo. In: Ueshima, R. (Ed.), Catalogue of invertebrate collection deposited in the Department of Zoology, the University Museum, the University of Tokyo (3). Material Reports No. 90. The University Museum, The University of Tokyo, Tokyo, pp. 1–57.

    Orensanz, J.M. (1990) The Eunicemorph polychaete annelids from Antarctic and Subantarctic Seas. With addenda to the Eunicemorpha of Argentina, Chile, New Zealand, Australia, and the Southern Indian Ocean. Antarctic Research Series, 52, 1–183.

    Parapar, J., Besteiro, C. & Urgorri, V. (1993) Taxonomy and Ecology of Annelida of the Iberian Peninsula—Polychaeta from the Ria-De-Ferrol. Cahiers de Biologie Marine, 34, 411–432.

    Pettibone, M.H. (1963) Marine polychaete worms of the New England region, 1 Aphroditidae through Trochochaetidae. Bulletin of the United States National Museum, 227, 1–356.

    https://doi.org/10.5479/si.03629236.227.1

    Salazar-Vallejo, S.I. (2014) Revision of Pherusa Oken, 1807 (Polychaeta: Flabelligeridae). Zootaxa, 3886 (1), 1–61.

    https://doi.org/10.11646/zootaxa.3886.1.1

    Salazar-Vallejo, S.I. & Carrera-Parra, L.F. (1998) Eunícidos (Polychaeta) del Caribe mexicano con claves para las especies del Gran Caribe: Fauchaldius, Lysidice, Marphysa, Nematonereis y Palola. Revista Biología Tropical, 45, 1481–1498.

    Schmarda, L.K. (1861) Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise un die Erdr 1853 bis 1857. Erster Band (zweite halfte) Turbellarian, Rotatorien un Anneliden, 2, 1–164.

    Steiner, T.M. & Amaral, A.C.Z. (2000) Two new species of Marphysa Quatrefages, 1865 (Eunicidae, Polychaeta) from intertidal sandy beaches of the Sao Sebastiao Channel, State of Sao Paulo (Brazil). Bulletin of Marine Science, 67 (1), 479–490.

    Treadwell, A.L. (1906) Polychaetous annelids of the Hawaiian Islands collected by the steamer Albatross in 1902. Bulletin of the United States Fish Commission, 23 (3), 1145–1181.

    Treadwell, A.L. (1917) A new species of polychaetous annelid from Panama, with notes on a Hawaiian form. Proceedings of the United States National Museum, 52 (2186), 427–430.

    https://doi.org/10.5479/si.00963801.52-2186.427

    Treadwell, A.L. (1921) Leodicidae of the West Indian region. Carnegie Institution of Washington Publication, 15 (293), 1–131.

    https://doi.org/10.5962/bhl.part.20335

    Treadwell, A.L. (1939) New polychaetous annelids from New England, Texas and Puerto Rico. American Museum Novitates, 1023, 1–7.

    Verrill, A.E. (1873) XVIII. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, with an account of the physical characters of the region. Report on the condition of the sea fisheries of the south coast of New England [later becomes Reports of the United States Commissioner of Fisheries], 1, 295–778.

    https://doi.org/10.5962/bhl.title.57652

    von Marenzeller, E. (1888) Polychäten der Angra Pequena-Bucht. Zoologische Jahrbücher, Abteilung für Systematik, Geographie und Biologie der Tiere, 3 (1), 1–24.

    Wang, Z., Zhang, Y. & Qiu, J.W. (2018) A new species in the Marphysa sanguinea Complex (Annelida, Eunicidae) from Hong Kong. Zological Studies, 57 (48), 1–13.

    https://doi.org/10.6620/ZS.2018.57-48

    Webster, H.E. (1879) The Annelida Chaetopoda of the Virginian coast. Transactions Albany Institute, 9, 202–269.

    https://doi.org/10.5962/bhl.title.11296

    Webster, H.E. (1884) Annelida from Bermuda collected by G. Brown Goode. Bulletin United States National Museum, 25, 307–327.

    Wijnhoven, S. & Dekker, A. (2010) Records of a new alien polychaete worm species, Marphysa sanguinea (Montagu, 1815) (Eunicidae) in the Eastern Scheldt, the Netherlands. Aquatic Invasions, 5 (4), 431–436.

    https://doi.org/10.3391/ai.2010.5.4.13

    Zanol, J., Halanych, K.M., Struck, T.H. & Fauchald, K. (2010) Phylogeny of the bristle worm family Eunicidae (Eunicida, Annelida) and the phylogenetic utility of noncongruent 16S, COI and 18S in combined analyses. Molecular Phylogenetics and Evolution, 55 (2), 660–676.

    https://doi.org/10.1016/j.ympev.2009.12.024

    Zanol, J., Halanych, K.M. & Fauchald, K. (2014) Reconciling taxonomy and phylogeny in the bristleworm family Eunicidae (Polychaete: Annelida). Zoologica Scripta, 43, 79–100.

    https://doi.org/10.1111/zsc.12034

    Zanol, J., da Silva, T.D.S.C. & Hutchings, P.A. (2016) Marphysa (Eunicidae, polychaete, Annelida) species of the sanguinea group from Australia, with comments on pseudo-cryptic species. Invertebrate Biology, 135, 328–344.

    https://doi.org/10.1111/ivb.12146