Skip to main content Skip to main navigation menu Skip to site footer
Published: 2021-07-30

Revision of the squat lobsters of the genus Phylladiorhynchus Baba, 1969 (Crustacea, Decapoda, Galatheidae) with the description of 41 new species

Centre d’Estudis Avançats de Blanes (CEAB-CSIC), C. acc. Cala Sant Francesc, 14, 17300 Blanes, Girona, Spain. Museo Nacional de Ciencias Naturales (MNCN-CSIC), José Gutiérrez Abascal, 2, 28006 Madrid, Spain.
Centre d’Estudis Avançats de Blanes (CEAB-CSIC), C. acc. Cala Sant Francesc, 14, 17300 Blanes, Girona, Spain.
Museo Nacional de Ciencias Naturales (MNCN-CSIC), José Gutiérrez Abascal, 2, 28006 Madrid, Spain.
Mitochondrial markers diversity taxonomy systematics Galatheoidea Anomura Decapoda Indian Ocean Pacific Ocean Crustacea


The genus Phylladiorhynchus Baba, 1969 currently contains 11 species, all occurring in the shallow waters and on the continental shelf of the Indian and Pacific oceans. Recent expeditions in these oceans have resulted in the collection of numerous new specimens in need of analysis. We have studied this material using an integrative approach analysing both morphological and molecular (COI and 16S) characters. We describe 41 new species and resurrect three old names: P. integrus (Benedict, 1902) and P. lenzi (Rathbun, 1907), previously synonymized with P. pusillus (Henderson, 1885), and P. serrirostris (Melin, 1939), previously synonymized with P. integrirostris (Dana, 1852). Most species of the genus are described and illustrated. Some species are barely discernible on the basis of morphological characters but are highly divergent genetically. Species of Phylladiorhynchus are mainly distinguishable by the number of epigastric spines and lateral spines of the carapace, the shape and the armature of the rostrum, the number and pattern of the ridges on the carapace and pleon, the shape of thoracic sternite 3 and the armature of the P2–4 dactyli. A dichotomous identification key to all species is provided.


  1. Ahyong, S.T. (2007) Decapod Crustacea collected by the NORFANZ expedition: Galatheidae and Polychelidae. Zootaxa, 1593, 1–54.

  2. Andrade, H. (1985) Crustáceos decápodos marinos del Archipiélago de Juan Fernandez. In: Arana, P. (Ed.), Investigaciones marinas en el Archipiélago de Juan Fernandez.Chile. Universidad Católica de Valparaíso, Valparaíso, pp. 109–116.

  3. Baba, K. (1969) Four new genera with their representatives and six new species of the Galatheidae in the collection of the Zoological Laboratory, Kyushu University, with redefinition of the genus Galathea. Ohmu, 2, 1–32.

  4. Baba, K. (1977) Biological results of the Snellius Expedition XXVIII. The galatheid Crustacea of the Snellius Expedition. Zoologische Mededelingen, Leiden, 50, 243–259.

  5. Baba, K. (1979) Expédition Rumphius II (1975) Crustacés parasites, commensaux, etc. (Th. Monod et R. Sèrene, éd.) VII. Galatheid crustaceans (Decapoda, Anomura). Bulletin du Muséum national d’Histoire naturelle, Paris, 4e Série, Section A, 1, 643–657.

  6. Baba, K. (1989) Anomuran crustaceans obtained by dredging from Oshima Strait, Amami-Oshima of the Ryukyu Islands. Memoirs of the National Science Museum, Tokyo, 22, 127–134.

  7. Baba, K. (1990) Chirostylid and galatheid crustaceans of Madagascar (Decapoda, Anomura). Bulletin du Muséum national d’Histoire naturelle, Paris, 4e Série, Section A, 11, 921–975.

  8. Baba, K. (1991) Crustacea Decapoda: Alainius gen. nov., Leiogalathea Baba, 1969, and Phylladiorhynchus Baba, 1969 (Galatheidae) from New Caledonia. In: Crosnier, A. (Ed.), Résultats des Campagnes MUSORSTOM. Vol. 9. Mémoires du Muséum national d’Histoire naturelle, Paris, (A), 152, 479–491.

  9. Baba, K. (1993) Anomoeomunida, a new genus proposed for Phylladiorhynchus caribensis Mayo, 1972 (Crustacea: Decapoda: Galatheidae). Proceedings of the Biological Society of Washington, 106, 102–105.

  10. Baba, K. (2005) Deep-sea chirostylid and galatheid crustaceans (Decapoda: Anomura) from the Indo-West Pacific, with a list of species. Galathea Reports, 20, 1–317.

  11. Baba, K., Macpherson, E., Poore, G.C.B., Ahyong, S.T., Bermudez, A., Cabezas, P., Lin, C.W., Nizinski, M., Rodrigues, C. & Schnabel, K.E. (2008) Catalogue of squat lobsters of the world (Crustacea: Decapoda: Anomura-families Chirostylidae, Galatheidae and Kiwaidae). Zootaxa, 1905 (1), 1–220.

  12. Baba, K., Macpherson, E., Lin, C.-W. & Chan, T.-Y. (2009) Crustacean fauna of Taiwan: squat lobsters (Chirostylidae and Galatheidae). National Taiwan Ocean University, Keelung, 312 pp.

  13. Baba, K., Ahyong, S.T. & Macpherson, E. (2011) Morphology of marine squat lobsters. In: Poore, G.C.B., Ahyong, S.T. & Taylor, J. (Eds.), The biology of squat lobsters. CSIRO Publishing, Melbourne, and CRC Press, Boca Raton, Florida, pp. 1–38.

  14. Balss, H. (1913) Neue Galatheiden aus der Ausbeute der deutschen Tiefsee-Expedition. ‘Valdivia’. Zoologischer Anzeiger, 41, 221–226.

  15. Balss, H. (1922) Decapoden von Juan Fernandez. In: Skottsberg, C. (Ed.), The Natural History of Juan Fernandez and Easter Island. Zoology, Uppsala, pp. 329–340.

  16. Benedict, J.E. (1902) Description of a new genus and forty-six new species of crustaceans of the family Galatheidae with a list of the known marine species. Proceedings of the Biological Society of Washington, 26, 243–334.

  17. Borradaile, L.A. (1916) Crustacea. Part 1.—Decapoda. British Antarctic (Terra Nova) Expedition, 1910. Natural History Reports Zoology, 3, 75–110, figs. 111–116.

  18. Cabezas, P., Macpherson, E. & Machordom, A. (2010) Taxonomic revision of the genus Paramunida Baba, 1988 (Crustacea: Decapoda: Galatheidae): a morphological and molecular approach. Zootaxa, 2712 (1), 1–60.

  19. Cabezas, P., Lin, C.W. & Chan, T.Y. (2011) Two new species of the deep-sea squat lobster genus Munida Leach, 1820 (Crustacea: Decapoda: Munididae) from Taiwan: morphological and molecular evidence. Zootaxa, 3036 (1), 26–38.

  20. Castilla, J.C. & Rozbaczylo, N. (1987) Invertebrados marinos de Isla de Pascua y Sala y Gómez. Islas oceánicas chilenas: conocimiento científico y necesidades de investigación. In: Castilla, J.C. (Ed.), Islas Oceánicas Chilenas: Conocimiento científicos y necesidades de investigaciones. Ediciones Univ Católica de Chile, Santiago, pp. 191–215

  21. Castro, P. (2011) Catalog of the anomuran and brachyuran crabs (Crustacea: Decapoda: Anomura, Brachyura) of the Hawaiian Islands. Zootaxa, 2947, 1–154.

  22. Chilton, C. (1906) Report of some Crustacea dredged off the coast of Auckland. Transactions and proceedings of the New Zealand Institute, 38, 265–269.

  23. Chilton, C. (1911) Crustacea. Scientific Results New Zealand Governm. Trawling Expedition 1907. Records of the Canterbury Museum, 1, 285–312.

  24. Crandall, K.A. & Fitzpatrick Jr, J.F. (1996) Crayfish molecular systematics: using a combination of procedures to estimate phylogeny. Systematic Biology, 45, 1–26.

  25. Dana, J.D. (1852) Crustacea. Part I. United States Exploring Expedition, during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U.S.N., 13, 1–685, with a folio atlas of 96 pls (published 1885).

  26. Davie, P.J.F. (2002) Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. CSIRO Publishing, Melbourne, xiv + 641 pp.

  27. De los Ríos Escalante, P. & Ibáñez Arancibia, E. (2016) A checklist of marine crustaceans known from Easter Island. Crustaceana, 89, 63–84.

  28. DiSalvo, L.H., Randall, J.E. & Cea, A. (1988) Ecological Reconnaissance of the Easter Island Sublittoral Marine-Environment. National Geographic Research, 4 (4), 451–473.

  29. Dong, C. & Li, X.Z. (2013) Galatheid squat lobster species from Chinese waters. Chinese Journal of Oceanology and Limnology, 31, 1315–1321.

  30. 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 (5), 294–9.

  31. Fujita, Y. (2007) First zoeas of two shallow-water galatheids, Lauriea gardineri (Laurie, 1926) and Phylladiorhynchus integrirostris (Dana, 1853) (Crustacea; Decapoda; Anomura; Galatheidae). Proceedings of the Biological Society of Washington, 120, 74–85.[74:FZOTSG]2.0.CO;2

  32. Grant, F.E. & McCulloch, A.R. (1906) On a collection of Crustacea from the Port Curtis district, Queensland. Proceedings of the Linnean Society of New South Wales, 1906, 2–53, pls. 1–4.

  33. Guiler, E.R. (1952) A list of the Crustacea of Tasmania. Records of the Queen Victoria Museum, Launceston, 3 (3), 15–44.

  34. Haig, J. (1955) Reports of the Lund University Chile Expedition 1948–49. 20. The Crustacea Anomura of Chile. Lunds Universtitets Arsskrift, Neue Folge, Avd., 2, 51, 1–68.

  35. Haig, J. (1973) Galatheidea (Crustacea, Decapoda, Anomura) collected by the F.I.S. Endeavour. Records of the Australian Museum, 28, 269–289.

  36. Hale, H.M. (1927) The Crustaceans of South Australia. Part I. Government Printer, Adelaide, 201 pp.

  37. Henderson, J.R. (1885) Diagnoses of new species of Galatheidae collected during the “Challenger” expedition. Annals and Magazine of Natural History, Series V, 16, 407–421.

  38. Henderson, J.R. (1888) Report on the Anomura collected by H.M.S. Challenger during the years 1873–76. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–76. Zoology, 27, 1–221, 21 pls.

  39. Katoh, K., Misawa, K., Kuma, K.I. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30 (14), 3059–3066.

  40. Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33 (7), 1870–1874.

  41. Larsson, A. (2014) AliView: a fast and lightweight alignment viewer and editor for large data sets. Bioinformatics, 30 (22), 3276–3278.

  42. Laurie, R.D. (1926) Reports of the Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the leadership of Mr. J. Stanley Gardiner, M.A. Vol. 8. No. VI. Anomura collected by Mr. J. Stanley Gardiner in the western Indian Ocean in H.M.S. Sealark. Transactions of the Linnean Society of London, Series 2, Zoology, 19, pp. 121–167, pls. 128 + 129.

  43. Leach, W.E. (1820) Galatéadées. In: Dictionnaire des Sciences Naturelles. F. G. Levreault, Paris, pp. 49–56.

  44. Lee, S.-H., Lee, S.-K., Kim, S.H. & Kim, W. (2019) First records of two squat lobsters (Decapoda, Galatheidae) from Korea. Crustaceana, 92, 725–737.

  45. Lenz, H. (1902) Die Crustaceen der Sammlung Plate (Decapoda und Stomatopoda). Zoologische Jahrbücher. Abteilung für Systematik, 5, 731–772, pl. 23.

  46. Lewinsohn, C. (1969) Die Anomuren des Roten Meeres (Crustacea Decapoda: Paguridea, Galatheidea, Hippidea). Zoologische Verhandelingen, Leiden, 104, 1–213, pls 1–2.

  47. Lewinsohn, C. (1982) Phylladiorhynchus integrirostris (Dana) und Lauriea gardineri (Laurie) (Decapoda, Anomura) aus dem nordlichen Roten Meer. Crustaceana, 42, 295–301.

  48. Machordom, A. & Macpherson, E. (2004) Rapid radiation and cryptic speciation in squat lobsters of the genus Munida (Crustacea, Decapoda) and related genera in the South West Pacific: molecular and morphological evidence. Molecular Phylogenetics and Evolution, 33, 259–279.

  49. Machordom, A., Araujo, R., Erpenbeck, D. & Ramos, M.Á. (2003) Phylogeography and conservation genetics of endangered European Margaritiferidae (Bivalvia: Unionoidea). Biological Journal of the Linnean Society, 78, 235–252.

  50. Macpherson, E. (2008) Some new records of shallow-water galatheid crustaceans (Anomura: Galatheidae) from the Dampier Archipelago, Western Australia. Records of the Western Australian Museum Supplement, 72, 289–297.

  51. Macpherson, E. & Baba, K. (2011) Taxonomy of squat lobsters. In: Poore, G.C.B., Ahyong, S.T. & Taylor, J. (Eds.), The biology of squat lobsters. CSIRO Publishing, Melbourne and CRC Press, Boca Raton, Florida, pp. 39–71.

  52. Macpherson, E. & Robainas-Barcia, A. (2013) A new genus and some new species of the genus Lauriea Baba, 1971 (Crustacea, Decapoda, Galatheidae) from the Pacific and Indian Oceans, using molecular and morphological characters. Zootaxa, 3599 (2), 136–160.

  53. Macpherson, E. & Robainas-Barcia, A. (2015) Species of the genus Galathea Fabricius, 1793 (Crustacea, Decapoda, Galatheidae) from the Indian and Pacific Oceans, with descriptions of 92 new species. Zootaxa, 3913 (1), 1–335.

  54. Macpherson, E., Rodríguez-Flores, P.C. & Machordom, A. (2020) Squat lobsters of the families Munididae and Munidopsidae from Papua New Guinea. In: Corbari, L. Ahyong, S.T. & Chan, T.-Y. (Eds.), Deep-sea crustaceans from Papua New Guinea. Tropical Deep-Sea Benthos 31. Mémoires du Muséum national d’Histoire naturelle, Paris, 213, pp. 11–120.

  55. Makarov, V.V. (1938) Anomura. Rakoobraznyey. Vol. 10. No. 3. In: Shtakel’berg, A.A. (Ed.), Fauna SSSR. New Series. Vol. 16. Academii Nauk SSSR, Moscow, pp. i–x + 1–324, 5 pls. [English translation, 1962: Crustacea, Anomura. Israel Program for Scientific Translation, Jerusalem, 278 pp.]

  56. Mayo, B. (1972) Three new species of the family Galatheidae (Crustacea, Anomura) from the western Atlantic. Bulletin of Marine Science, 22, 522–535.

  57. Melin, G. (1939) Paguriden und Galatheiden von Prof. Dr. Sixten Bocks Expedition nach den Bonin-Inseln 1914. Kungliga Svenska Vetenskapsakademiens Handlingar, 18, 1–119.

  58. Meusnier, I., Singer, G.A.C., Landry, J.-F., Hickey, D.A., Hebert, P.D.N. & Hajibabaei, M. (2008) A universal DNA mini-barcode for biodiversity analysis. BioMed Central Genomics, 9, 214.

  59. Miyake, S. (1965) Crustacea, Anomura. In: Okada, Y.K. & Uchida, T. (Eds.), New illustrated encyclopedia of the fauna of Japan. s.n, Tokyo, pp. 630–652.

  60. Miyake, S. & Baba, K. (1965) Some galatheids obtained from the Bonin Islands (Crustacea, Anomura). Journal of the Faculty of Agriculture, Kyushu University, 13, 585–593.

  61. Miyake, S. & Baba, K. (1966) Descriptions of galatheids collected from coral reefs of the Ryukyu Islands (Crustacea, Anomura) Journal of the Faculty of Agriculture, Kyushu University, 14, 57–79.

  62. Miyake, S. & Baba, K. (1967) Galatheids of the East China Sea (Chirostylidae and Galatheidae, Decapoda, Crustacea) Journal of the Faculty of Agriculture, Kyushu University, 14, 225–246.

  63. Miyake, S. & Nakazawa, K. (1947) Crustacea, Anomura. In: Uchida, S. (Ed.), Illustrated encyclopedia of the fauna of Japan (exclusive of insects). Revised edition. Hokuryukan, Tokyo, pp. 731–750.

  64. Mujica, A., González-Cornejo, F., Meerhoff, E. & Yannicelli, B. (2019) Larval development of Phylladiorhynchus pusillus (Henderson, 1885) (Decapoda, Anomura, Galatheidae). Latin American Journal of Aquatic Research, 47, 774–783.

  65. Palumbi, S., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) Simple fool’s guide to PCR. University of Hawaii, Honolulu, 45 pp.

  66. Peyrot-Clausade, M. (1989) Crab cryptofauna (Brachyura and Anomura) of Tikehau, Tuamotu Archipelago, French Polynesia. Coral Reefs, 8, 109–117.

  67. Poore, G.C.B. (2004) Marine decapod Crustacea of southern Australia. A guide to identification (with chapter on Stomatopoda by Shane Ahyong). CSIRO Publishing, Melbourne, 574 pp.

  68. Poore, G.C.B., McCallum, A. & Taylor, A.C. (2008) Decapod Crustacea of the continental margin of southwestern adn central Western Australia: preliminary identifications of 524 species from FRV Southern Surveyor voyage SS10-2005. Museum Victoria Science Report, 11, 1–106.

  69. Porter, C.E. (1916a) Materiales para la fauna carcinologica de Chile. XII.—Sobre los Galatheidae del Museo Nacional. Revista Chilena de Historia Natural, 20, 111–117.

  70. Porter, C.E. (1916b) Los crustaceos decapodos Chilenos del Museo Nacional (estudios criticos) III.—La fam. Galatheidae. Boletin Museo Nacional de Historia Natural, Chile, 9, 95–100.

  71. Poupin, J. (1996) Crustacea Decapoda of French Polynesia (Astacidea, Palinuridea, Anomura, Brachyura). Atoll Research Bulletin, 442, 1–114.

  72. Poupin, J. (2003) Crustacea Decapoda and Stomapoda of Easter Island and surrounding areas. A documented checklist with historical overview and biogeographic comments. Atoll Research Bulletin, 500, 1–50.

  73. Rathbun, M.J. (1907) South American Crustacea. Revista Chilena de Historia Natural, 11, 45–50, fig. 41, pls. 42 + 43.

  74. Retamal, M.A. (1981) Catálogo ilustrado de los crustáceos decápodos de Chile. Gayana Zoología, 44, 1–110.

  75. Retamal, M.A. (2004) Decápodos de las islas oceánicas chilenas: Pascua y Salas y Gómez. Ciencia y Tecnología del Mar, 27, 55–68.

  76. Retamal, M.A. & Arana, P.M. (2016) Record of stomatopods and decapods, including descriptions of the species of commercial interest from the submarine rises and surrounding waters of the Chilean oceanic islands (southeastern Pacific Ocean). Latin American Journal of Aquatic Research, 44, 16–33.

  77. Richer de Forges, B., Chan, T.Y., Corbari, L., Lemaitre, R., Macpherson, E., Ahyong, S.T. & Ng, P.K.L. (2013) The MUSORSTOM-TDSB deep-sea benthos exploration program (1976–2012): An overview of crustacean discoveries and new perspectives on deep-sea zoology and biogeography. In: Ahyong, S.T., Chan, T.-Y., Corbari, L. & Ng, P.K.L. (Eds.), Tropical Deep-Sea Benthos 27. Mémoires du Muséum national d’Histoire naturelle, 204, pp. 13–66.

  78. Rodríguez-Flores, P.C., Machordom, A. & Macpherson, E. (2017) Three new species of squat lobsters of the genus Fennerogalathea Baba, 1988 (Decapoda: Galatheidae) from the Pacific Ocean. Zootaxa, 4276 (1), 46–60.

  79. Rodríguez-Flores, P.C., Macpherson, E., Buckley, D. & Machordom, A. (2019a) High morphological similarity coupled with high genetic differentiation in new sympatric species of coral-reef squat lobsters (Crustacea, Decapoda, Galatheidae). Zoological Journal of the Linnean Society, 185, 984–1017.

  80. Rodríguez-Flores, P.C., Macpherson, E. & Machordom, A. (2019b) Revision of the squat lobsters of the genus Leiogalathea Baba, 1969 (Crustacea, Decapoda, Munidopsidae) with the description of 15 new species. Zootaxa, 4560 (2), 201–256.

  81. Rodríguez-Flores, P.C., Machordom, A., Abelló, P., Cuesta, J.A. & Macpherson, E. (2019c). Species delimitation and multi-locus species tree solve an old taxonomic problem for European squat lobsters of the genus Munida Leach, 1820. Marine Biodiversity, 49, 1751–1773.

  82. Rowden, A.A., Schnabel, K.E., Schlacher, T.A., Macpherson, E., Ahyong, S.T. & Richer de Forges, B. (2010) Squat lobster assemblages on seamounts differ from some, but not all, deep-sea habitats of comparable depth. Marine Ecology, 31, 63–83.

  83. Samouelle, G. (1819) The entomologists’ useful compendium; or an introduction to the knowledge of British Insects, comprising the best means of obtaining and preserving them, and a description of the apparatus generally used; together with the genera of Linné, and modern methods of arranging the Classes Crustacea, Myriapoda, spiders, mites and insects, from their affinities and structure, according to the views of Dr. Leach. Also an explanation of the terms used in entomology; a calendar of the times of appearance and usual situations of near 3,000 species of British Insects; with instructions for collecting and fitting up objects for the microscope. Thomas Boys, London, 496 pp.

  84. Schnabel, K.E. & Ahyong, S.T. (2019) The squat lobster genus Phylladiorhynchus Baba, 1969 in New Zealand and eastern Australia, with description of six new species. Zootaxa, 4688 (3), 301–347.

  85. Thomson, G.M. (1899) A revision of the Crustacea Anomura of New Zealand. Transactions of the Royal Society of New Zealand, Zoology, 31, 169–197.

  86. Tirmizi, N.M. (1966) Crustacea: Galatheidae. The John Murray Expedition 1933–34. Scientific Reports, 11, 167–234.

  87. Tirmizi, N.M. & Javed, W. (1980) Two new species and one new record of Phylladiorhynchus Baba from the Indian Ocean (Decapoda, Galatheidae) Crustaceana, 39, 255–262.

  88. Tirmizi, N.M. & Javed, W. (1993) Indian Ocean galatheids (Crustacea: Anomura) Marine Reference Collection and Resource Centre, University of Karachi, Karachi, 147 pp

  89. Whitelegge, T. (1900) Crustacea. Part 1. Scientific results of the trawling expedition of H.M.C.S. “Thetis”, off the coast of New South Wales, in February and March, 1898. Australian Museum Memoir, 4, 135–199, pls. 32–35.

  90. Yokoya, Y. (1936) Some rare and new species of decapod crustaceans found in the vicinity of the Misaki Marine Biological Station. Japanese Journal of Zoology, 7, 129–146.

  91. Zarenkov, N.A. (1968) Crustacea Decapoda collected in the Antarctic and Antiboreal regions by the Soviet Antarctic Expedition. Issledovaniya Fauny Morei, SSSR, 6, 153–199. [in Russian]