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Issue: Vol. 5249 No. 5: 6 Mar. 2023
Type: Article
Published: 2023-03-06
DOI: 10.11646/zootaxa.5249.5.2
Page range: 540-558
Abstract views: 395
PDF downloaded: 47

Recognition of Parapinnixidae Števčić, 2005, and Tetriasidae fam. nov. in Pinnotheroidea De Haan, 1833 (Crustacea: Brachyura: Thoracotremata)

Simon F. S. Li Marine Science Laboratory, School of Life Sciences, The Chinese University of Hong Kong, Shatin, Hong Kong, China.
Tropical Biosphere Research Center, Iriomote Station, University of the Ryukyus, 870 Uehara, Taketomi, Okinawa 907–1541, Japan
Tetriasidae new family Parapinnixidae pea crab Pinnotheridae Pinnotheroidea phylogenetics systematics Crustacea

Abstract

The systematic position of three aberrant pinnotheroid genera: Tetrias Rathbun, 1898, Parapinnixa Holmes, 1895 and Sakaina Serène, 1964, are reappraised. A new family, Tetriasidae fam. nov. is established for Tetrias, and Parapinnixidae Števčić, 2005, is recognised for Parapinnixa and Sakaina. Tetriasids differ from all other pinnotheroids in that the articles of the palp of the third maxilliped (carpus, propodus and dactylus) are large and the dactylus is inserted at the distolateral corner of the propodus, whereas parapinnixids are distinctive in having the buccal cavern relatively small, subtriangular with the third maxilliped ischiomerus small, mesioproximally produced to form a triangular projection and is completely fused. The two families are phylogenetically distinct from the family Pinnotheridae based on the multi-locus gene tree, and are basal or near basal lineages in the Pinnotheroidea.

References

  1. Ahyong, S.T. (2018) Revision of Ostracotheres H. Milne Edwards, 1853 (Crustacea: Brachyura: Pinnotheridae). Raffles Bulletin of Zoology, 66, 538–571.

  2. Ahyong, S.T. & Ng, P.K.L. (2007) The pinnotherid type material of Semper (1880), Nauck (1880) and Bürger (1895) (Crustacea: Decapoda: Brachyura). Raffles Bulletin of Zoology, Supplement 16, 191–226.

  3. Ahyong, S.T. & Ng, P.K.L. (2009) Aphanodactylidae, a new family of thoracotreme crabs (Crustacea: Brachyura) symbiotic with polychaete worms. Zootaxa, 2289 (1), 33–47. https://doi.org/10.11646/zootaxa.2289.1.3

  4. Alcock, A. (1900) Materials for a carcinological fauna of India. No. 6. The Brachyura Catometopa, or Grapsoidea. Journal of the Asiatic Society of Bengal, 69 (2), 279–456. https://doi.org/10.5962/bhl.title.15344

  5. Becker, C. & Türkay, M. (2010) Taxonomy and morphology of European pea crabs (Crustacea: Brachyura: Pinnotheridae). Journal of Natural History, 44 (25–26), 1555–1575. https://doi.org/10.1080/00222931003760020

  6. Bouvier, E.-L. (1917) Gonoplacidés et Pinnothéridés nouveaux recueillis au cours des campagnes américaines du “Hassler” et du “Blake”. Bulletin du Muséum National d’Histoire Naturelle, Paris, 23 (5), 391–398.

  7. Bürger, O. (1895) Ein Beitrag zur Kenntniss der Pinnotherinen. Zoologischen Jahrbüchern, 8, 361–390.

  8. Campos, E. (1994) Parapinnixa cubana, a new pea crab from Cuba (Crustacea: Brachyura: Pinnotheridae). Proceedings of the Biological Society of Washington, 107 (2), 308–311.

  9. Campos, E. (2009) A new species and two new genera of pinnotherid crabs from the northeastern Pacific Ocean, with a reappraisal of the subfamily Pinnotherinae de Haan, 1833 (Crustacea: Brachyura: Pinnotheridae). Zootaxa, 2022 (1), 29–44. https://doi.org/10.11646/zootaxa.2022.1.3

  10. Campos, E. (2017) Taxonomy of Pinnotheres bipunctatus Nicolet, 1849 with a distributional checklist of the Pinnotheridae of Chile and Peru, and a list of the Crustacea described by Hercule Nicolet in the Atlas of the Physical and Political History of Chile. Latin American Journal of Aquatic Research, 45 (2), 379–390. https://doi.org/10.3856/vol45-issue2-fulltext-13

  11. Campos, E. & Griffith, H. (1990) Clypeasterophilus, a new genus to receive the small-palped species of the Dissodactylus complex (Brachyura: Pinnotheridae). Journal of Crustacean Biology, 10 (3), 550–553. https://doi.org/10.2307/1548346

  12. Campos, E. & Wicksten, M.K. (1997) A new genus for the Central American crab Pinnixa costaricana Wicksten, 1982 (Crustacea: Brachyura: Pinnotheridae). Proceedings of the Biological Society of Washington, 110 (1), 69–73.

  13. Castro, P. (2015) Symbiotic Brachyura. In: Castro, P., Davie, P.J.F., Guinor, D., Schram, F.R., von Vaupel Klein, J.C. (Eds.), Treatise on Zoology—Anatomy, Taxonomy, Biology. The Crustacea. Vol. 9 Part C-I. Brill, Leiden, pp. 543–581. https://doi.org/10.1163/9789004190832_012

  14. Colgan, D., McLauchlan, A., Wilson, G. & Livingston, S. (1998) Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology, 46, 419–437. https://doi.org/10.1071/ZO98048

  15. Crandall, K.A. & Fitzpatrick Jr., J.F. (1996) Crayfish molecular systematics: using a combination of procedures to estimate phylogeny. Systematic Biology, 45, 1–26. https://doi.org/10.1093/sysbio/45.1.1

  16. Cuesta, J.A., Schubart, C.D. & Felder, D.L. (2005) Systematic position of the Asthenognathinae Stimpson, 1858 and Pseudopinnixa carinata Ortmann (Decapoda, Brachyura) new findings from larval and DNA comparisons. In: The Sixth International Crustacean Congress. Glasgow, July 2005, pp. 127. [abstract]

  17. Dana, J.D. (1851) Conspectus Crustaceorum quæ in Orbis Terrarum circumnavigatione, Carolo Wilkes e Classe Reipublicæ Fœderatæ Duce, lexit et descripsit. Proceedings of the Academy of Natural Sciences of Philadelphia, 5, 247–254.

  18. Davie, P.J.F. & Ng, N.K. (2007) A new genus for cave-dwelling crabs previously assigned to Sesarmoides (Crustacea: Decapoda: Brachyura: Sesarmidae). Raffles Bulletin of Zoology, Supplement 16, 227–231.

  19. De Haan, H.M. (1833–1850) Crustacea. In: von Siebold, P.F., Fauna Japonica, sive Descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batavia imperium tenent, suscepto, annis 1823–1830. Apud Auctorem, Lugduni Batavorum, pp. xxx + i–xxxi + vii–xvii + 243 + viii–xvi pp., pls. I–LII + 2 + A–Q.

  20. De Gier, W. & Becker, C. (2020) A review of the ecomorphology of pinnotherine pea crabs (Brachyura: Pinnotheridae), with an updated list of symbiont-host associations. Diversity, 12 (11), 431. https://doi.org/10.3390/d12110431

  21. Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797. https://doi.org/10.1093/nar/gkh340

  22. Garth, J.S. (1957) Reports of the Lund University Chile Expedition 1948–49. 29. The Crustacea Decapoda Brachyura of Chile. Lunds Universitets Årsskrift, Ny fölid, Avdelningen 2, 53 (7), 1–129.

  23. Glassell, S.A. (1933) Notes on Parapinnixa affinis Holmes and its allies. Transactions of the San Diego Society of Natural History, 7 (27), 319–330. https://doi.org/10.5962/bhl.part.6163

  24. Glassell, S.A. (1938) New and obscure decapod Crustacea from the west American coasts. Transactions of the San Diego Society of Natural History, 8 (33), 411–454.

  25. Guinot, D. (1977) Propositions pour une nouvelle classification des Crustacés Décapodes Brachyoures. Comptes rendus hebdomadaires des séances de l’Académie des sciences, Série D, 285, 1049–1052.

  26. Green, T.M. (1992) Pinnaxodes gigas, a new species of pinnotherid crab from the Gulf of California (Decapoda: Brachyura: Pinnotheridae) Proceedings of the Biological Society of Washington, 105 (4), 775–779.

  27. Heard, R.W. & Manning, R.B. (1997) Austinixa, a new genus of pinnotherid crab (Crustacea: Decapoda: Brachyura), with the description of A. hardyi, a new species from Tobago, West Indies. Proceedings of the Biological Society of Washington, 110 (3), 393–398.

  28. Holmes, S.J. (1895) Notes on west American Crustacea. Proceedings of the California Academy of Sciences, series 2, 4, 563–588, pls. 20–21. [Dated 1894, published 1895]

  29. Holmes, S.J. (1900) Synopsis of California stalk-eyed Crustacea. Occasional Papers of the California Academy of Sciences, 7, 1–263. https://doi.org/10.5962/bhl.title.53705

  30. Holthuis, L.B. (1975) Limotheres, a new genus of pinnotherid crab, commensal of the bivalve Lima, from the Caribbean Sea. Zoologische Mededelingen, Leiden, 48 (25), 291–295.

  31. Hultgren, K.M., Foxx, C.L. & Palacios Theil, E. (2022) Host‑associated morphological convergence in symbiotic pea crabs. Evolutionary Ecology, 36, 272–286. https://doi.org/10.1007/s10682-022-10153-0

  32. Jiang, W. & Liu, R (2011) New species and new records of pinnotherid crabs (Crustacea: Decapoda: Brachyura) from the Yellow Sea. Zoologischer Anzeiger, 250, 488–496. https://doi.org/10.1016/j.jcz.2011.05.001

  33. Kim, H.S. & Sakai, T. (1972) A new pinnotherid crab of the genus Sakaina (decapod crustaceans) from Korea. Proceeding of the Japanese Society of Systematic Zoology, 8, 29–31.

  34. Kobjakova, Z.I. (1967) Decapoda (Crustacea, Decapoda) from the Possjet Bay (the Sea of Japan). Explorations of the Fauna of the Seas of the USSR, 5 (13), 220–248. [in Russian]

  35. Komai, T. & Konishi, K. (2012) Reappraisal of the Systematic Position of the Supposed Pinnotherid Crab Pseudopinnixa carinata (Crustacea: Decapoda: Brachyura). Species Diversity, 17, 29–37. https://doi.org/10.12782/sd.17.1.029

  36. Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2017) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34, 772–773. https://doi.org/10.1093/molbev/msw260

  37. Liao, Y., De Grave, S., Ho, T.W., Ip, B.H.Y., Tsang, L.M., Chan, T.Y. & Chu, K.H. (2017) Molecular phylogeny of Pasiphaeidae (Crustacea, Decapoda, Caridea) reveals systematic incongruence of the current classification. Molecular Phylogenetic Evolution, 115, 171–180. https://doi.org/10.1016/j.ympev.2017.07.021

  38. Linnaeus, C. (1767) s.n. In: Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Editio duodecima, reformata. Regnum Animale. Tom. I. Pars II. Vol. 1. Laurentii Salvii, Holmiae, pp. 533–1327. https://doi.org/10.5962/bhl.title.156772

  39. Lockington, W.N. (1877) Remarks on the Crustacea of the Pacific coast of North America, including a catalog of the species in the Museum of the California Academy of Sciences, San Francisco. Proceedings of the California Academy of Sciences, 7, 63–78. [for 1876] https://doi.org/10.5962/bhl.title.37872

  40. MacLeay, W.S. (1838) On the brachyurous decapod Crustacea brought from the Cape by Dr. Smith. In: Smith, A. (Ed.), Illustrations of the Annulosa of South Africa; being a portion of the objects of natural history chiefly collected during an expedition into the interior of South Africa, under the direction of Dr. Andrew Smith, in the years 1834, 1835. and 1836; fitted out by “The Cape of Good Hope Association for Exploring Central Africa”. Smith, Elder, and Co., London, pp. 53–71, pls. 2 + 3. https://doi.org/10.1080/00222933809512348

  41. Manning, R.B. (1993) West African pinnotherid crabs, subfamily Pinnotherinae (Crustacea, Decapoda, Brachyura). Bulletin du Muséum national d’Histoire naturelle, Section A, Zoologie, Biologie et Ecologie Animales, Paris, 4e série, 15 (1–4), 125–177.

  42. Manning, R.B. & Morton, B. (1987) Pinnotherids (Crustacea: Decapoda) and Leptonaceans (Mollusca: Bivalvia) associated with sipunculan worms in Hong Kong. Proceedings of the Biological Society of Washington, 100 (3), 543–551.

  43. Marques, F. & Pohle, G. (1995) Phylogenetic analysis of the Pinnotheridae (Crustacea, Brachyura) based on larval morphology, with emphasis on the Dissodactylus species complex. Zoolgica Scripta, 24, 347–364. https://doi.org/10.1111/j.1463-6409.1995.tb00479.x

  44. Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. 2010 gateway computing environments workshop (GCE), 2010, 1–8. https://doi.org/10.1109/GCE.2010.5676129

  45. Milne-Edwards, A. (1867) Descriptions de quelques espèces nouvelles de Crustacés Brachyures. Annales de la Société Entomologique de France, Serie 4, 7, 263–288.

  46. Milne-Edwards, A. (1873) Recherches sur la faune carcinologique de la Nouvelle-Calédonie, II. Nouvelles Archives du Muséum d’Histoire naturelle, Paris, 9, 155–332, pls. 4–18. https://doi.org/10.5962/bhl.title.10410

  47. Milne Edwards, H. (1852) De la famille des ocypodides (Ocypodidae). Second Mémoire. Observations sur les affinités zoologiques et la classification naturelle des crustacés. Annales des Sciences Naturelles, 3e Série, 18, 128–166, pls. 3–4.

  48. Milne Edwards, H. (1853) Mémoire sur la famille des Ocypodides. Suite (1). Deuxiéme Tribu Principale. Annales des Sciences Naturelles, 3e Série, 20, 163–228, pls. 6–11.

  49. Milne Edwards, H. & Lucas, H. (1842–1844) Crustacés. In: d’Orbigny, A. (Ed.), Voyage dans l’Amerique méridionale (le Brésil, la république orientale de l’Uruguay, la république Argentine, la Patagonie, la république du Chili, la république de Bolivia, la république du Pérou), exécutée pendant les années 1826, 1827, 1828, 1829, 1830, 1831, 1832 et 1833. Vol. 6 (1). P. Bertrand, Vve Levrault, Paris and Strasbourg, pp. 1–37, +1–17.

  50. Minh, B.Q., Nguyen, M.A.T. & von Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30, 1188–1195. https://doi.org/10.1093/molbev/mst024

  51. Nauck, E. (1880) Das Kaugerüst der Brachyuren. Zeitschrift für wissenschaftliche Zoologie, Leipzig, 34, 1–69, pl. I, 2 woodcuts.

  52. Ng, P.K.L. (2018) On the identities of Pinnotheres villosissimus Doflein, 1904, P. dofleini Lenz, in Lenz & Strunck, 1914, and P. pilumnoides Nobili, 1906 (Decapoda, Brachyura, Pinnotheridae) from the western Indian Ocean. Crustaceana, 91 (5), 611–633 https://doi.org/10.1163/15685403-00003785

  53. Ng, P.K.L. & Ahyong, S.T. (2022) The pea crab genus Arcotheres Manning, 1993 (Crustacea: Brachyura: Pinnotheridae) from Singapore and Peninsular Malaysia, with a reappraisal of diagnostic characters and descriptions of two new genera. Raffles Bulletin of Zoology, 70, 134–248.

  54. Ng, P.K.L. & Ho, P.-H. (2016) A new genus for Fabia obtusidentata Dai, Feng, Song and Chen, 1980, a pea crab (Decapoda: Brachyura: Pinnotheridae) symbiotic with the Moon Scallop Amusium pleuronectes (Linnaeus, 1758) (Mollusca: Pectinidae). Journal of Crustacean Biology, 36 (5), 740–751. https://doi.org/10.1163/1937240X-00002474

  55. Ng, P.K.L., Ahyong, S.T. & Campos, E. (2019) Two new genera of pinnotherid crabs (Crustacea: Brachyura: Pinnotheroidea) from the Americas and the Western Pacific. Raffles Bulletin of Zoology, 67, 337–351.

  56. Ng, P.K.L., Guinot, D. & Davie, P.J.F. (2008) Systema Brachyurorum: part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology, Supplement 17, 1–286.

  57. Ng, P.K.L. & Manning, R.B. (2003) On two new genera of pea crabs parasitic in holothurians (Crustacea: Decapoda: Brachyura: Pinnotheridae) from the Indo-West Pacific, with notes on allied genera. Proceedings of the Biological Society of Washington, 116 (4), 901–919.

  58. Nguyen, L., 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

  59. Ortmann, A.E. (1894) Die Decapoden-Krebse des Strassburger Museums, mit besonderer Berücksichtigung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit im Strassburger Museum aufbewahrten Formen. VIII Thiel. Abtheilung: Brachyura (Brachyura Genuina Boas), III. Unterabtheilung: Cancroidea, 2. Section: Cancrinea, 2. Gruppe: Catametopa. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Thiere, 7 (5), 683–772, pl. 23. https://doi.org/10.5962/bhl.part.24064

  60. Palacios Theil, E. & Felder, D.L. (2020) Phylogeny of the genus Pinnixa White, 1846 (Crustacea, Brachyura, Pinnotheridae) and allies inferred from mitochondrial and nuclear molecular markers, with generic reassignment of twenty-one species. Zoosystema, 42 (6), 85­­–103. https://doi.org/10.5252/zoosystema2020v42a6

  61. Palacios-Theil, E., Cuesta, J.A., Campos, E. & Felder, D.L. (2009) Molecular genetic re-examination of subfamilies and polyphyly in the family Pinnotheridae (Crustacea: Decapoda). In: Martin, J.W., Crandall, K.A. & Felder, D.L. (Eds.), Crustacean Issues 18: Decapod Crustacean Phylogenetics. CRC Press, Taylor & Francis Group, Boca Raton, Florida, pp. 457–474.

  62. https://doi.org/10.1201/9781420092592-c23

  63. Palacios Theil, E., Cuesta, J.A. & Felder, D.L. (2016) Molecular evidence for non-monophyly of the pinnotheroid crabs (Crustacea: Brachyura: Pinnotheroidea), warranting taxonomic reappraisal. Invertebrate Systematics, 30, 1–27. https://doi.org/10.1071/IS15023

  64. Pohle, G. & Marques, F. (1998) Phylogeny of the Pinnotheridae: larval and adult evidence, with emphasis on the evolution of gills. Invertebrate Reproduction & Development, 33, 229–239. https://doi.org/10.1080/07924259.1998.9652635

  65. Poore, G.C.B. & Ahyong, S.T. (2023) Marine decapod Crustacea: a guide to the families and genera of the world. CSIRO Publishing, Collingwood, 916 pp. https://doi.org/10.1071/9781486311798

  66. Rathbun, M.J. (1898) The Brachyura collected by the U.S. Fish Commission steamer Albatross on the voyage from Norfolk, Virginia, to San Francisco, California, 1887–1888. Proceedings of the United States National Museum, 21, 567–616, pls. 41–44. https://doi.org/10.5479/si.00963801.21-1162.567

  67. Rathbun, M.J. (1918) The grapsoid crabs of America. Bulletin of the United States National Museum, 97, 1–461, pls. 1–161. https://doi.org/10.5479/si.03629236.97.i

  68. Rathbun, M.J. (1931) A new species of pinnotherid crab from Costa Rica. Journal of the Washington Academy of Sciences, 21 (12), 262–263.

  69. Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542. https://doi.org/10.1093/sysbio/sys029

  70. Sakai, T. (1969) Two new genera and twenty-two new species of crabs from Japan. Proceedings of the Biological Society of Washington, 82, 243–280.

  71. Say, T. (1817–1818) An account of the Crustacea of the United States. Journal of the Academy of Natural Sciences, Philadelphia, 1, 57–63 + 65–80 (pl. 4) + 97–101 + 155–160 + 161–169 + 235–253 + 313–319 + 374–380 + 381–401 + 423–441.

  72. Schmitt, W.L., McCain, J.C. & Davidson, E.S. (1973) Decapoda I. Brachyura I. Family Pinnotheridae. Crustaceorum Catalogus. Vol. 3. W. Junk B.V., The Hague, 160 pp.

  73. Serène, R. (1964) Goneplacidae et Pinnotheridae. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914–1916, 80, 181–282, pls. 16–24.

  74. Simon, C., Frati,F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 651–701. https://doi.org/10.1093/aesa/87.6.651

  75. Smith, S.I. (1869) Notice of the Crustacea collected by Prof. C. F. Hartt on the coast of Brazil in 1867. Transactions of the Connecticut Academy of Arts and Sciences, 2, 1–41, pl. I. https://doi.org/10.5962/bhl.part.20634

  76. Števčić, Z. (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica, 14, 1–159.

  77. Stimpson, W. (1858) Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et descripsit. Pars V. Crustacea Ocypodoidea. Proceedings of the Academy of Natural Sciences of Philadelphia, 10, 93–110. https://doi.org/10.5962/bhl.title.51447

  78. Takeda, M. & Prince Masahito (2000) Systematic notes on the pinnotherid crabs of the genus Pinnaxodes (Crustacea: Decapoda: Brachyura). Bulletin of the National Science Museum, Tokyo, Series A, 26 (3), 99–112.

  79. Takeda, M., Maruyama, S. & Konishi, K. (1991) Taxonomical and biogeographical notes on Sakaina yokoyai (Glassell) from northern Japan (Crustacea: Decapoda: Pinnotheridae). Bulletin of the Biogeographical Society of Japan, 46 (9), 89–95.

  80. Thoma, B.P., Heard, R.W. & Vargas, R. (2005) A new species of Parapinnixa (Decapoda: Brachyura: Pinnotheridae) from Isla del Coco, Costa Rica. Proceedings of the Biological Society of Washington, 118 (3), 543–550. https://doi.org/10.2988/0006-324X(2005)118[543:ANSOPD]2.0.CO;2

  81. Tsang, L.M., Ma, K.Y., Ahyong, S.T., Chan, T.Y. & Chu, K.H. (2008) Phylogeny of Decapoda using two nuclear protein-coding genes: origin and evolution of the Reptantia. Molecular Phylogenetics Evolution, 48, 359–368. https://doi.org/10.1016/j.ympev.2008.04.009

  82. Tsang, L.M., Chan, B.K.K., Shih, F.L., Chu, K.H. & Chen, A.C.L. (2009) Host associated speciation in the coral barnacle Wanella milleporae (Cirripedia: Pygomatidae) inhabiting the Millepora coral. Molecular Ecology, 18, 1463–1475. https://doi.org/10.1111/j.1365-294X.2009.04090.x

  83. Tsang, L.M., Chan, T.Y., Ahyong, S.T. & Chu, K.H. (2011) Hermit to king, or hermit to all: multiple transitions to crab-like forms from hermit crab ancestors. Systematic Biology, 60, 616–629. https://doi.org/10.1093/sysbio/syr063

  84. Tsang, L.M., Schubart, C.D., Ahyong, S.T., Lai, J.C.Y., Au, E.Y.C., Chan, T.Y., Ng, P.K.L. & Chu, K.H. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and the origin of freshwater crabs. Molecular Biology and Evolution, 31, 1173–1187. https://doi.org/10.1093/molbev/msu068

  85. Tsang, L.M., Ahyong, S.T., Shih, H.T. & Ng, P.K.L. (2018) Further polyphyly of pinnotheroid crabs: the molecular phylogenetic position of the polychaete-associated Aphanodactylidae. Invertebrate Systematics, 32, 92–99. https://doi.org/10.1071/IS17038

  86. Tsang, C.T.T., Schubart, C.D.S., Chu, K.H., Ng, P.K.L. & Tsang, L.M. (2022) Molecular phylogeny of Thoracotremata crabs (Decapoda, Brachyura): toward adopting monophyletic superfamilies, invasion history into terrestrial habitats and multiple origins of symbiosis. Molecular Phylogenetics and Evolution, 177, 107596. https://doi.org/10.1016/j.ympev.2022.107596

  87. Wells, H.W. & Wells, M.J. (1961) Observations on Pinnaxodes floridensis, a new species of pinnotherid crustacean commensal in holothurians. Bulletin of Marine Science of the Gulf and Caribbean, 11 (2), 267–279.

  88. Werding, B. & Müller, H.G. (1990) Die Gattung Parapinnixa Holmes 1894 an der Nordküste Kolumbiens, mit Beschreibung von Parapinnixa magdalenensis n. sp. (Crustacea: Decapoda: Pinnotheridae). Senckenbergiana biologica, 70 (1/3), 221–227.

  89. White, A. (1846) Notes on four new genera of Crustacea. Annals and Magazine of Natural History, Series 1, 18, 176–178, pl. 2. https://doi.org/10.1080/037454809494406

  90. Wicksten, M.K. (1982) Pinnixa costaricana, a new species of crab from Central America (Brachyura: Pinnotheridae). Proceedings of the Biological Society of Washington, 95, 579–582.