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The non-indigenous bryozoan Amathia verticillata (delle Chiaje, 1822) is recorded for the first time from the southern Mexican Pacific. The species was detected between 2014 and 2018 at three locations along the coast of Oaxaca, southwestern Mexico: Puerto Ángel Bay, Santa Cruz Bay, and La Blanca Island. A discussion on the introduction of the species to Oaxaca is also included.

Hirose, M., Gordon, D.P. (2020): New seriated Amathia species in Japan, with a redescription of A. acervata Lamouroux, 1824 (Bryozoa: Ctenostomata). Zootaxa 4742 (2): 311-331, DOI: https://doi.org/10.11646/zootaxa.4742.2.5

Amathia fimbria n. sp. (Figs 4, 5) Material examined. Holotype: NSMT-Bry R55, single bushy colony on polychaete tube, Najima, Sagami Bay, Honshu, no depth data, 10 June 1934. Paratypes: NSMT-Bry R235, colony on hydroid, Udorishima, Kasajima, Sagami Bay, Honshu, 9 m, 26 July 1964; NSMT-Bry R375, two bushy colonies on hydroid, off Miihama, Suzaki, Izu, Honshu, 10–20 m, 22 June 1983; NSMT-Bry R381, bushy colonies on hydroid stem, Ooura, Suzaki, Izu, Honshu, 10 m, 19 June 1987. Other material examined: NSMT-Bry R 179, dried specimen with small colony, South Hirashima, Mitsuiso, off Nagaoka, Kaneda Bay, Honshu, 14 m, 15 July 1962. Etymology. The specific name derives from the Latin fimbria which means fringe or tassel, referring to the compact ball-shaped colony morphology resembling Kikutoji which is a kind of tassel for Japanese traditional clothes Kimono. Description. Colony (Fig. 4A, B) erect, bushy and densely branching; initially tufted, comprising stolons attached to a substratum, from which erect stems arise at intervals. As colonies grow, cross connections can occur so that the lower half of a colony tends to appear as a tangle, with branching in all planes. Colonies anchored by rhizoid-like stolons adherent to the substratum, becoming thick-walled with age. Tips of branches can also adhere to the substratum, continuing as adherent stolons, producing, at intervals, 5 opposing pairs of non-connate autozooids, each autozooid at an oblique angle to the stolon and somewhat flattened against the substratum, the whole cluster appearing overall like a pinna (Fig. 4E). The largest colony up to 60 mm high and 70 mm across, anchored by a short (Remarks. This species is similar to the holotype of A. acervata and A. brevisilva n. sp. described above; it differs in having trichotomous branching of the stems; it also has occasional pinnate branching of adherent parts of stolons. In this latter character, Amathia fimbria n. sp. is similar to Amathia reptopinnata n. sp. described below, however it differs from the latter species in having a shorter internode stem and shorter autozooid cluster with less spirality. The present species also resembles Amathia rudis Kubanin, 1992 in modes of branching and some other features. The present species, however, differs from Kubanin’s (1992) description of A. rudis in having shorter and narrower stolons and a slightly higher value of the proportion of the autozooid cluster on the stolon. The present species also differs from A. rudis in having pinnate autozooid clusters adhering to the substratum branch tips. The material in the Showa Collection occurs on a polychaete tube, hydroid stems and an undetermined species of Sargassum. Distribution. Southern Sagami Bay region, the Pacific coast of Honshu, Japan, 9–20 m deep., Published as part of Hirose, M. & Gordon, D. P., 2020, New seriated Amathia species in Japan, with a redescription of A. acervata Lamouroux, 1824 (Bryozoa: Ctenostomata), pp. 311-331 in Zootaxa 4742 (2) on pages 320-322, DOI: 10.11646/zootaxa.4742.2.5, http://zenodo.org/record/3677823, {"references":["Kubanin, A. (1992) New bryozoan species (Gymnolaemata, Ctenostomata) from the Japan Sea. Zoologcheskii Zhurnal, 71, 19 - 31. [in Russian]"]}

Amathia reptopinnata n. sp. (Figs 6–8) ? Amathia distans: Mawatari, 1948: p. 11; 1963: p. 6. Material examined. Holotype: NSMT-Te1213, colony on leaves of eelgrass (Zostera sp.), Lake Komuke, Hokkaido, 44°15’58.36” N, 143°30’53.70” E, ~ 1 m, 10 August 2007. Paratypes: NSMT-Te893, 894, colonies on leaves of eelgrass (Zostera sp.), same collection data as for holotype, Lake Komuke, Hokkaido, 44°15’58.36” N, 143°30’53.70” E, ~ 1 m, 10 August 2007; NSMT-Te895–897, colonies on leaves of eelgrass (Zostera sp.), Lake Saroma, Hokkaido, 44°11’7.00” N, 143°45’48.60” E, ~ 1 m, 6 October 2010; NSMT-Te898–900, some branches of colonies, entrance to Kamo Bay, Oki Islands, Shimane Prefecture, Honshu, 36°10’23.77” N, 133°16’50.04” E– 36°10’28.08” N, 133°16’48.72” E, 13–16 m, 26 June 2012; NSMT-Te901, colonies on oyster shell, collected at Matsushima Bay, Miyagi Prefecture, 2 m in depth, 11 November 2013. Etymology. The specific name derives from the Latin reptilis (creeping) and pinnatus (pinnate), referring to the pinnately arranged, prostrate zooids. Description. Colony (Fig. 6A) erect, tufted, fairly densely branching, up to 50 mm high, comprising stolons attached to a substratum, from which erect stems arise at intervals. As colonies grow, occasional cross connections can occur (Fig. 7A, B). Colonies anchored by rhizoidal stolons adherent to the substratum, becoming thick-walled with age (Fig. 7C). Tips of branches can also adhere to the substratum, continuing as adherent stolons with short, planar grappling-hook-like processes at intervals along their length or with blind side branches also with a planar, grappling-hook morphology (Fig. 7C, E). Adherent stolons also producing, at intervals, up to 11 opposing pairs of non-connate autozooids, each autozooid at an oblique angle to the stolon and flattened against the substratum, the whole cluster appearing overall like a pinna (Fig. 7D, F). Erect growths are initiated as one main stem with side branches, and branching is always dichotomous (Fig. 6B, C). Stolon segments more or less straight or gently curving, with a mean length of 2.60± 0.62 mm (range 1.62–3.99 mm, n = 86), the angle between branches at dichotomies 45–80°. Stolon with white or light-brown pigments (Fig. 6C). Mean stolon width 0.18± 0.04 mm (range 0.12–0.26 mm, n = 89). Autozooid clusters (Fig. 8) disposed in clockwise spirals on most stolon segments (some segments with anticlockwise spirals and others lacking a cluster) in Matsushima, whereas mostly anticlockwise in Oki, and clockwise or anticlockwise depending on the colony in Hokkaido. Autozooid clusters comprising 12–23 ‘pairs’ per cluster, each cluster describing at least one complete 360° circuit of the stolon from its commencement to its completion, or up to 2.3 turns; mean cluster length 1.45± 0.21 mm (range 1.01–1.98 mm, n = 99); each cluster has a mean linear distance of 1.21± 0.43 mm (range 0.57–2.23 mm, n = 79) on the stolon and terminates at the next branch node or at the next stolon septum (there can be three stolon segments between branch nodes); zooid cluster occupying 39–76% of stolon length. Autozooids tilted distad c. 60° from the perpendicular, with a mean length of 0.45± 0.05 mm (range 0.36–0.53 mm, n = 48), connate; zooid width (as measured in lateral view of zooid cluster) averaging 0.14± 0.02 mm (range 0.11–0.19 mm, n = 47); owing to the zooid tilt, the distal end of each cluster generally leans past the point of termination of the zooid insertions at each branch node. Outer walls of zooids slightly thicker than interior walls, evidenced by darker cuticularisation, and the outer distal rim of the cuticle can be slightly flared. Tentacle number 8. Descending rhizoids evident in proximal parts of larger, older colonies, each appearing to issue from a point immediately proximal to an autozooid cluster. Remarks. This species is similar to the previous species but is slightly more robust, darker in colour, and with dichotomous branching instead of being mostly trichotomous. The mean and maximum lengths of stolons and autozooid clusters are also larger than the previous species. The present species resembles Amathia brasiliensis Busk, 1886, Amathia vidovici Heller, 1867, and Amathia aegyptiana d'Hondt, 1983 in that neighboring autozooids are connate for half their length. Amathia reptopinnata n. sp. differs, however, from A. brasiliensis, A. vidovici, and A. distans in having a greater maximum number of zooid pairs per autozooid cluster. Amathia reptopinnata n. sp. differs from A. aegyptiana in having a slightly narrower stolon, and pinnately arranged autozooid clusters at attachment areas on the substratum. Amathia reptopinnata n. sp. resembles Amathia similis Gordon & Spencer Jones, 2013 in the dimensions of colony, stolon and autozooid, but differs by having both a larger proportion of occupancy of the autozooid cluster on the stolon and a pinnately arranged autozooid cluster on the substratum. Although the present specimen from Matsushima is similar to A. rudis in stolon length, it differs by having a typically larger proportion of occupancy of the autozooid cluster on the stolon; A. reptopinnata n. sp. from Matsushima (and Oki and Hokkaido) also has a longer autozooid cluster and pinnately arranged prostrate zooids. The present species differs from A. acervata in having longer stolon segments and longer autozooid clusters. Colonies grow on the leaves of eelgrass (Zostera sp.) in Lakes Saroma and Komuke (northern Hokkaido), on oyster shells in Matsushima Bay (northeastern Honshu) and on Sargassum sp. at Oki Islands (Japan Sea), at depths of 1–16 m depending on locality and substratum. Distribution. Japan Sea off western Honshu, the Pacific coast of Tohoku region, and northern Hokkaido in the Okhotsk Sea., Published as part of Hirose, M. & Gordon, D. P., 2020, New seriated Amathia species in Japan, with a redescription of A. acervata Lamouroux, 1824 (Bryozoa: Ctenostomata), pp. 311-331 in Zootaxa 4742 (2) on pages 322-326, DOI: 10.11646/zootaxa.4742.2.5, http://zenodo.org/record/3677823, {"references":["Mawatari, S. (1948) Bryozoan fauna of Matsushima Bay, northern Honshu, Japan (preliminary report). Proceedings of the Biogeographical Society of Japan, 2, 1 - 5.","Busk, G. (1886) Report on the Polyzoa collected by H. M. S. Challenger during the years 1873 - 76. Part II. - The Cyclostomata, Ctenostomata, and Pedicellinea. Report on the Scientific Results of the Voyage of H. M. S. Challenger, Zoology, 17 (3), 50, i-viii + 1 - 47, 10 pls.","Heller, C. (1867) Die Bryozoen des adriatischen Meeres. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 17, 77 - 136.","Gordon, D. P. & Spencer Jones, M. E. (2013) The amathiiform Ctenostomata (phylum Bryozoa) of New Zealand-including four new species, two of them of probable alien origin. Zootaxa, 3647 (1), 75 - 95. https: // doi. org / 10.11646 / zootaxa. 3647.1.4"]}

Amathia brevisilva n. sp. (Fig. 3) ? Amathia distans: Okada & Mawatari, 1938: p. 446; Mawatari, 1952: p. 262. ? Bowerbankia gracilis: Mawatari, 1952: p. 262. Material examined. Holotype: NSMT-Te1209, single colony on pebble, Manazuru, Sagami Bay, 35° 9'23.78"N, 139° 8'38.56"E, intertidal (~ 1 m), 7 August 2017. Paratypes: NSMT-Te1210, Te1211, some branches, same collection data as for holotype, Manazuru, Sagami Bay, 35° 9'23.78"N, 139° 8'38.56"E, intertidal (~ 1 m), 7 August 2017; SMBL-30 and NSMT-Te1212, colony on hydroid stem, single specimen with several labels in Seto Marine Biological Station, Kyoto University and a small fragments in NSMT, collected at Kitahama and/or off Tonda, Wakayama Prefecture, no depth data, collected by beach seine, 15–21 April 1928 and/or 13 August 1929 and/or March 1944, possibly the specimens reported in the study by Okada and Mawatari (1938) and/or Mawatari (1952). Etymology. The specific name derives from the Latin brevis (small/low) and silva (bush, forest, or mass), referring to the small bushy colony morphology growing on the surface of the substrate. Description. Colony (Fig. 3A, B) erect, small, sparsely branching; narrow stolons attached to a substratum, from which erect stems arise at intervals. As colonies grow, cross connections can occur so that the lower half of a colony tends to appear as a tangle, with branching in all planes. Colonies anchored by narrow rhizoid-like stolons adherent to the substratum; lacking any autozooids on the substratum. The largest colony up to 40 mm high and 55 mm across, anchored by a short (Remarks. This species is very similar to the holotype of A. acervata described above in the short stolon length, width and zooid-cluster length; it differs in having fewer autozooid pairs per cluster. In having a short stolon length, Amathia brevisilva n. sp. is similar to Amathia fimbria n. sp. described below, but differs from the latter species in having dichotomous branches. The present species also resembles Amathia rudis Kubanin, 1992 in the spirality of the autozooid cluster and some other features. The holotype colony of A. rudis was collected from the southwestern part of Peter the Great Gulf in the Japan Sea, in the Posyet area at a depth of 2 m on blades of Zostera; paratype material was found a little further to the east in Novgorod Bay at 3 m depth on a pier. Among the most characteristic features noted by Kubanin was the branching, comprising variable dichotomy, trichotomy and even a rare tetrachotomy. He described the colony as creeping, but attached mainly in the area of the ancestrula and rarely elsewhere. The form of the spiral of autozooidal clusters on stolon segments was not described but his figures seem to indicate both anticlockwise and clockwise spirality. The polypide had eight tentacles. The present species, however, differs from Kubanin’s (1992) description of A. rudis in having narrower and shorter stolons and slightly shorter autozooid clusters. This species had been reported as Amathia gracilis and/or Amathia imbricata in various previous reports (Mawatari 1948, 1952, 1962); however, it differs from A. gracilis in erect branching colony morphology and from A. imbricata in less density of zooids along the stolon. This is very likely the species that Okada & Mawatari (1938) and Mawatari (1952) reported as Amathia distans, Bowerbankia [i.e. Amathia] imbricata, and Bowerbankia [i.e. Amathia] gracilis from various localities in Wakayama Prefecture, including Kii Peninsula. Distribution. Sagami Bay and Kii Peninsula, the Pacific coast of Honshu, Japan, from the intertidal to the shallow nearshore subtidal, on pebbles and hydroid stems., Published as part of Hirose, M. & Gordon, D. P., 2020, New seriated Amathia species in Japan, with a redescription of A. acervata Lamouroux, 1824 (Bryozoa: Ctenostomata), pp. 311-331 in Zootaxa 4742 (2) on pages 317-320, DOI: 10.11646/zootaxa.4742.2.5, http://zenodo.org/record/3677823, {"references":["Okada, Y. & Mawatari, S. (1938) On the collection of Bryozoa along the coast of Wakayama-ken, the middle part of Honsyu, Japan. Annotationes Zoologicae Japonensis, 17, 445 - 463.","Mawatari, S. (1952) Bryozoa of the Kii Peninsula. Publications of the Seto Marine Biological Laboratory, 2, 261 - 288, pl. 12. https: // doi. org / 10.5134 / 174675","Kubanin, A. (1992) New bryozoan species (Gymnolaemata, Ctenostomata) from the Japan Sea. Zoologcheskii Zhurnal, 71, 19 - 31. [in Russian]","Mawatari, S. (1948) Bryozoan fauna of Matsushima Bay, northern Honshu, Japan (preliminary report). Proceedings of the Biogeographical Society of Japan, 2, 1 - 5.","Mawatari, S. (1962) Bryozoa of the eastern shore of Noto Peninsula. Annual report of the Noto Marine Laboratory / Noto Marine Laboratory, Faculty of Science, University of Kanazawa, 3, 5 - 10."]}

Amathia acervata Lamouroux, 1824 (Fig. 2) Amathia acervata Lamouroux, 1824: p. 45; Jelly, 1899: p. 10; d’Hondt, 1991: p. 165 (part). Serialaria acervata: Blainville, 1834: p. 476; Deshayes & Milne Edwards, 1836: p. 170; d’Orbigny, 1853: p. 595. Non Amathia acervata: d’Hondt, 1979: 10, 16; d’Hondt, 1983: p. 65, fig. 36E; d’Hondt, 1991: p. 163, 165 (part); Gordon et al., 2009: p. 288. [These all refer to Amathia bicornis Tenison-Woods, 1880.] Amathia vidovici: d’Hondt, 1991: p. 165 Material examined. Holotype: Muséum national d’Histoire naturelle (MNHN), Herbarium Sheet 86 (dried specimen; Fig. 2A), with original handwritten label (Fig. 2B); a reconstituted fragment of the holotype, now in 70% ethanol, accompanies the herbarium sheet. Collected in Japanese waters by Wilhelm Gottlieb Tilesius (during the first Russian circumnavigation on the Nadezhda), October 1804 (see Barratt 1981). Description (of reconstituted sample from holotype). Colonies erect, comprising relatively short (c. 12.0– 12.5 mm), sparsely branching tufts from small attachment points on algal substratum, these a little broader than the stolon and apparently somewhat triangular in cross section or roundly so; rhizoids not evident. Branching dichotomous (Fig. 2 C–F), at angles of 20–45°. Stolon segments more or less straight or very gently curving, sinuous where the autozooid pairs are clustered (Fig. 2E); mean stolon length 1.74± 0.38 mm (range 1.41–2.35 mm, n = 5). Mean stolon width 0.10± 0.01 mm (range 0.09–0.13 mm, n = 11). Stolon with refringent yellow granules (Fig. 2D). Autozooid clusters disposed in anticlockwise spirals on the stolon segments, comprising c. 12–15 ‘pairs’ per cluster, each cluster describing c. 1.6 turns about the stolon from its commencement to its completion; mean cluster length on stolon 1.00± 0.18 mm (range 0.75–1.29 mm, n = 8), terminating at the next stolon septum or branch node; mean inception distance of proximalmost zooid in cluster from bifurcation 0.82± 0.10 mm (range 0.66–1.00 mm, n = 9), zooid cluster occupying 54–75% of stolon length. Autozooids tilted distad from the perpendicular (Fig. 2 C–F), with a mean length of 0.44± 0.03 mm (range 0.37–0.48 mm, n = 9) in reconstituted dried retracted specimens, connate, the outermost half of each zooid wall with thicker cuticle (hemispherical when viewed apically or in optical section); zooid width (as measured in lateral view of zooid cluster) averaging 0.12± 0.01 mm (range 0.11–0.14 mm, n = 8); owing to the zooid tilt, the distal end of each cluster generally leans past the point of termination of the zooid insertions at each branch node. Remarks. Lamouroux (1824, p. 45) gave a diagnosis: “ Amathia dwarf, little branched, subdichotomous; hairlike branches, thin; subseparated cells (i.e. zooids) in distinct groups, together standing apart” (authors’ translation from the French). The fuller description of A. acervata given by Deshayes & Milne Edwards (1836) was translated by Tenison-Woods (1880) from the French thus: “the groups of cells are distant a millimetre from one another. They are composed of nearly 20 cells, heaped without order around the subdichotomous stems, which are not much branched, and isolated during the greater part of their length.” Not enough material is available (or able to be reconstituted) to determine the full range of characters or dimensions based on the type. There is considerable concordance or overlap of characters (see Table 2) between A. acervata and Amathia brevisilva n. sp., though there are some differences, such as the direction of spiral of the autozooid clusters. Lamouroux (1824) stated that his material came from the Sea of Japan (in French as “la mer du Japon ”) and that it had been collected by Wilhelm Gottlieb Tilesius (who visited Japan in 1804); subsequent French authors accepted this information (Blainville 1834; Deshayes & Milne Edwards 1836; d’Orbigny 1853). In recent decades, owing to a misunderstanding based on historic specimens in European museums, the binominal A. acervata was misapplied. d’Hondt (1979, 1983, 1991) inadvertently equated the species with Australian A. bicornis Tenison-Woods, 1880, treating the latter as a junior subjective synonym. However, the earliest descriptions of A. acervata sensu Lamouroux do not match published descriptions and illustrations of A. bicornis, which is a highly distinctive species. Lamouroux (1824) described the autozooids as heaped/stacked/accumulated without order, which is not at all true of the Australian taxon, which has highly ordered, connate biserial clusters of autozooids that spiral distinctively for 1.5 turns about the stolon segment. Each autozooid also carries a conspicuous pair of spine-like processes, hence the epithet bicornis; these are not mentioned in the diagnosis of A. acervata sensu Lamouroux. Neither specimen name (acervata or bicornis) is given in any of the comprehensive published 20th-century lists of Bryozoa from Japan (e.g. Mawatari 1965; Mawatari 1986) or anywhere else in East Asian seas subsequently (e.g. Liu et al. 2001; Seo 2005, 2011), until Seo et al. (2018) reported it from Korea’s ‘South Sea’ coast (based on advice from the present authors). Harmer (1915), however, had tentatively included A. acervata in the synonymy of putative A. distans, noting that, “It is not impossible that the present species is identical with A. acervata, described by Lamouroux (1824) from Japan.” d’Hondt (1979, p. 16) came to his conclusion concerning the synonymy of A. acervata and A. bicornis based on the examination of a specimen labelled ‘Sirinx spinosa’ by Charles-Alexandre Lesueur, according it the corrected name of “ Amathia acervata (Lamouroux, 1816 [sic]) (Holotype)” and “Nouvelle- Hollande?” as the provenance. However, he did not examine a Lamouroux specimen from Japan at that time as Lamouroux’s type was then thought lost (or destroyed in the Second World War). Subsequently, d’Hondt (1991) reported on a collection of Lamouroux Bryozoa rediscovered in the herbarium at the Université de Caen. The dried bryozoans were kept between dust covers in the manner of dried plant or algal specimens. Dust cover 86 pertained to Amathia acervata (Fig. 2A) retaining an original label from the time of Lamouroux (Fig. 2B); the specimen is clearly not like A. bicornis. Assuming A. acervata had been lost, d’Hondt (1991, p. 165) interpreted this material as belonging to a completely different species, Amathia vidovici (Heller, 1867). He incorrectly determined another specimen LBIMM-BRY- 4531 in the same collection that looks like A. bicornis as the syntype of A. acervata. In the information accompanying his original description, Lamouroux (1824) noted that A. acervata is ‘parasitic’ on Fucaceae. There is only one species of Fucus in Japan, viz. F. distichus subsp. evanescens, which occurs on the coast of Hokkaido, where Amathia has not been recorded in the published literature (see Mawatari 1972; Mawatari & Mawatari 1981). On the other hand, there are numerous species of Sargassum (Fucales, Sargassaceae) in Japan. Based on the discrepancies between the original descriptions and geographical distributions of A. acervata and A. bicornis, we conclude: 1) the specimen in dust-cover 86 labelled Amathia acervata and resembling A. vidovici (which would be not dissimilar from A. distans or look-alikes in the dried state) was in fact genuine A. acervata from Japan (redescribed in this study); 2) a specimen in the same collection (LBIMM-BRY-4531) that looks like A. bicornis came not from Japan but from Australia (from Tasmania via a Mr Cummings). Based on the dried holotype, d’Hondt (1991) noted that the European species Amathia vidovici (Heller, 1867) resembles A. acervata. Now that the characters of the latter have been clarified based on the reconstituted fragment, it is possible to make a more precise comparison: A. vidovici differs in having both clockwise and anticlockwise spirals that complete only a single turn about the stolon owing to fewer autozooid pairs in each cluster (8–9) and both Prenant & Bobin (1966) and Hayward & McKinney (2002) state that stolon diameter is 0.2 mm, rather larger than in any of the Japanese specimens discussed in this paper (see Table 2)., Published as part of Hirose, M. & Gordon, D. P., 2020, New seriated Amathia species in Japan, with a redescription of A. acervata Lamouroux, 1824 (Bryozoa: Ctenostomata), pp. 311-331 in Zootaxa 4742 (2) on pages 314-317, DOI: 10.11646/zootaxa.4742.2.5, http://zenodo.org/record/3677823, {"references":["Lamouroux, J. V. F. (1824) Amathie. In: Lamouroux, J. V. F., Bory de Saint Vincent, J. B. & Eudes-Deslongchamps, J. A. (Eds.), Encyclopedie d'Histoire naturelle des Zoophytes ou Animaux rayonees. Tome 95. Agasse, Paris, pp. 42 - 45.","Blainville, H. M. D. de (1834) Manuel d'Actinologie ou de Zoophytology. F. G. Levrault, Paris, 644 pp. https: // doi. org / 10.5962 / bhl. title. 8768","Deshayes, G. P. & Milne Edwards, H. (1836) Deuxieme Edition. Revue et augmentee de notes presentant les faits nouveaux dont la science s'est enrichie jusqu'a ce jour. In: Lamarck, J. B. P. A. de (Ed.), Histoire naturelle des Animaux sans Vertebres … precede d'une introduction offrant la determination des caracteres essentiels de l'animal, sa distinction du vegetal et des autres corps naturels, enfin, l'exposition des principes fondamentaux de la zoologie. Tome Deuxieme. Histoire des Polypes. J. B. Baillere, Paris, 684 pp.","Gordon, D. P., Taylor, P. D. & Bigey, F. P. (2009) Phylum Bryozoa - moss animals, sea mats, lace corals. In: Gordon, D. P. (Ed.), New Zealand Inventory of Biodiversity. Vol. 1. Radiata, Lophotrochozoa, Deuterostomia. Canterbury University Press, Christchurch, pp. 271 - 297.","Tenison-Woods, J. E. (1880) On the genus Amathia of Lamouroux, with a description of a new species. Transactions of the Royal Society of Victoria, 16, 89 - 118, 1 pl.","Barratt, G. R. V. (1981) Russia in Pacific Waters, 1715 - 1825: A survey of the origins of Russia's naval presence in the North and South Pacific. University of British Columbia Press, Vancouver, xvii + 300 pp.","Mawatari, S. (1965) Bryozoa. In: Okada, Y., Uchida, S. & Uchida, T. (Eds.), New Illustrated Encyclopedia of the Fauna of Japan. Hokuryu-Kan Publishing Co. Ltd, Tokyo, pp. 585 - 626. [in Japanese]","Mawatari, S. F. (1986) Bryozoans. In: The Japan Research Group of Marine Fouling (Ed. Comp.), Studies of Fouling Organisms. Koseisha-Koseikaku, Tokyo, pp. 71 - 106. [in Japanese]","Liu, X., Yin, X. & Ma, J. (2001) Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing, xxi + 860 pp., 82 pls. [in Chinese with English summary]","Seo, J. E. (2005) Bryozoa. Illustrated Encyclopedia of Fauna and Flora of Korea, 40, 1 - 596. [in Korean]","Seo, J. E. (2011) Bryozoa: Phylactolaemata, Stenolaemata, Gymnolaemata: Ctenostomata, Cheilostomata: Ascophora II. - Bryozoans. Invertebrate Fauna of Korea, 29, 1 - 107.","Seo, J. E., Chae, H. S., Winston, J. E., Zagorsek, K. & Gordon, D. P. (2018) Korean ctenostome bryozoans-observations on living colonies, new records, five new species, and an updated checklist. Zootaxa, 4486 (3), 251 - 283. https: // doi. org / 10.11646 / zootaxa. 4486.3.3","Harmer, S. F. (1915) The Polyzoa of the Siboga Expedition. Part 1. Entoprocta, Ctenostomata and Cyclostomata. Siboga-Expeditie, 28 a, 1 - 180, pls. 1 - 12.","Lamouroux, J. V. F. (1816) Histoire des Polypiers coralligenes flexibles, vulgairement nommes Zoophytes. Poisson, Caen, 560 pp. https: // doi. org / 10.5962 / bhl. title. 11172","Heller, C. (1867) Die Bryozoen des adriatischen Meeres. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 17, 77 - 136.","Mawatari, S. F. (1972) A new species of the genus Bowerbankia (Bryozoa, Ctenostomata) from Hokkaido. Journal of the Faculty of Science of Hokkaido University, Series 6, Zoology, 18, 300 - 304.","Mawatari, S. F. & Mawatari, S. (1981) A preliminary list of cheilostomatous bryozoans collected along the coast of Hokkaido. Proceedings of the Japanese Society of Systematic Zoology, 21, 41 - 58.","Hayward, P. J. & McKinney, F. K. (2002) Northern Adriatic Bryozoa from the vicinity of Rovinj, Croatia. Bulletin of the American Museum of Natural History, 270, 1 - 139. https: // doi. org / 10.1206 / 0003 - 0090 (2002) 270 % 3 C 0001: NABFTV % 3 E 2.0. CO; 2"]}

The invasive bryozoan Amathia verticillata has been associated with native biodiversity and economic losses around the world and in 2009, it was detected on one island of the Azores Archipelago and has since spread to other islands. The present study provides insights into the reproductive strategies of this species through the analysis of brooded embryos, the number of internodes and the degree of colony branching over a one year period in more than 200 colonies. These remote oceanic islands provide good environmental conditions for A. verticillata to diversify its gene pool, since the species is able to carry out sexual reproduction year-round. In contrast, population dynamics of A. verticillata seem to be strongly correlated with changes in temperature and the local photoperiod. Field intervention aimed at reducing the population in affected areas should be conducted in the winter when the number of potential colonizing fragments that might be generated by intervention is smaller.

The bryozoan Amathia (= Zoobotryon) verticillata was found near the northern entrance to the Port of Kaohsiung in November 2015 and is the first record from Taiwan. This bryozoan can form extensive colonies and is considered to be one of the more invasive nonindigenous species, occurring worldwide from Mediterranean to tropical environments within the Atlantic and Indo-Pacific Oceans. Colonies extending to approximately 80 cm in length were on immersed ropes attached to quay walls. It is likely that the species is established in the Port of Kaohsiung. Its distribution within the Pacific is reviewed.

Amathia cf. gracilis (Leidy, 1855) (Fig. 17) Bowerbankia gracilis: Seo 2011: 33, fig. 11 (cum syn., part). Material examined. MBRBK 1707. Seongsan port, Jeju Island. 33.4690° N, 126.9260° E, intertidal rocks, 8 May 2017. Description. Colonies in this species group consist of a network of stolons with erect, tubular and distally and proximally tapering zooids budded from them at intervals (Fig. 17A, C). Thriving colonies often grow into tangled masses, clumps and tufts. Zooids elongate-oval sacs, rounded at base and distally, with squared to puckered round orifice when polypide retracted. Retracted and preserved zooids c. 0.400 mm long, 0.060 mm wide, attached singly or in clusters on narrow creeping stolon. Internal organs and musculature seen through transparent walls of freshly budded zooids, but older zooids often fouled by algae or sediment particles. Polypide with conspicuous rounded gizzard and tentacle crown of 8 evenly spaced tentacles (Fig. 17A, B). No reproduction noted in colonies collected in this study. Remarks. Bowerbankia gracilis was originally described by Joseph Leidy (1855) from Rhode Island, USA, on the Western Atlantic seaboard where it is tolerant of reduced salinities and variations in temperature, occurring in estuaries and harbors and among fouling communities. It has been reported from many localities around the world, but some records may refer to other similar species. It has been reported from China (e.g. Liu et al. 2001), Japan, and Korea (Seo 2011). Distribution. Members of species group found almost worldwide in fouling and shallow water communities., Published as part of Seo, Ji-Eun, Chae, Hyun Sook, Winston, Judith E., Zágoršek, Kamil & Gordon, Dennis P., 2018, Korean ctenostome bryozoans-observations on living colonies, new records, five new species, and an updated checklist, pp. 251-283 in Zootaxa 4486 (3) on page 275, DOI: 10.11646/zootaxa.4486.3.3, http://zenodo.org/record/1453233, {"references":["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 Science Philadelphia, 3 (2), 135 - 152.","Seo, J. E. (2011) Bryozoa: Phylactolaemata, Stenolaemata, Gymnolaemata: Ctenostomata, Cheilostomata: Ascophora II. - Bryozoans. Invertebrate Fauna of Korea, 29, 1 - 107.","Liu, X., Yin & Ma, J. (2001) Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing, 860 pp., 82 pls."]}

Amathia acervata Lamouroux, 1824 (Fig. 16) Amathia acervata Lamouroux, 1824: 45; Jelly 1889: 10; Hirose et al. 2017. Serialaria acervata: Blainville 1834: 476; Deshayes & Milne Edwards 1836: 170; d’Orbigny 1853: 595. Non Amathia acervata: d’Hondt 1979: 10, 16; 1983: 65, fig. 36E; d’Hondt 1991: 163, 165 (part); Gordon et al. 2009: 288. [These all refer to A. bicornis.] Non Amathia vidovici: d’Hondt 1991: 165 [These refer to A. acervata]. Material examined. MBRBK1710, Tongyeong yacht anchorage, South Sea, 34.4934°N, 128.2600°E, 17 August 2015, depth 30 cm. Description. Colonies forming erect, bushy and branching tufts attached to algae and other substrata (Fig. 16A–D). Anticlockwise (usually) spirals of zooid clusters grouped along narrow (about 1.4 mm width) cylindrical stolons. Zooids tilted relative to stolon axis, narrow (about 0.5 mm long, 0.12 mm wide), tubular and connate and more thickly cuticularized at distal ends, a cluster consisting of 12–15 pairs (Fig. 16A, B). Clusters spiral about 1.5 turns around stolon (clusters c. 0.6–1.4 mm long). Stolon branching mostly trifurcate. Color of alcohol-preserved specimens whitish tan. Polypides with 8 tentacles. Remarks. Bushy Amathia species found in Asian waters have usually been attributed to Amathia convoluta (Lamarck, 1816) or Amathia distans Busk, 1886, species with Atlantic distributions. Hirose et al. (in prep.) have made a morphological and molecular study of three Amathia species in Japan, including the redescription of the holotype of Amathia acervata. This is the first record of the species in Korea. Another Japanese species, presently undescribed (Hirose et al. in prep.) is more robust, with slightly larger stolons and zooids with darker-brown pigmentation and mostly bifurcate branching. It may also be found to occur in Korea. No living specimens were found during our survey, but MBRBK houses jars of preserved material. Distribution. Korea: Japan Sea (East Sea) coastal localities. Also Pacific coast of Japan., Published as part of Seo, Ji-Eun, Chae, Hyun Sook, Winston, Judith E., Zágoršek, Kamil & Gordon, Dennis P., 2018, Korean ctenostome bryozoans-observations on living colonies, new records, five new species, and an updated checklist, pp. 251-283 in Zootaxa 4486 (3) on pages 274-275, DOI: 10.11646/zootaxa.4486.3.3, http://zenodo.org/record/1453233, {"references":["Lamouroux, J. V. F. (1824) Amathie. Encyclopedie d'Histoire Naturelle des Zoophytes ou Animaux rayonnes, 95, 42 - 45.","Jelly, E. C. (1889) A Synonymic Catalogue of Recent Marine Bryozoa. London, Dulau & Co., xv + 322 pp.","Hirose, M. (2017) Diversity of freshwater and marine bryozoans in Japan. In: Motokawa M. & Kajihara H. (Eds) Species Diversity of Animals in Japan. Diversity and Commonality in Animals. Springer, Tokyo, pp. 629 - 649.","Blainville, H. M. D. de (1834) Manuel d'Actinologie ou de Zoophytology. Paris, F. G. Levrault, 644 pp.","Deshayes, G. P. & Milne Edwards, H. (1836) Deuxieme Edition. Revue et augmentee de notes presentant les faits nouveaux dont la science s'est enrichie jusqu'a ce jour. In: Lamarck, J. B. P. A. de (Ed.), Histoire naturelle des Animaux sans Vertebres … precede d'une introduction offrant la determination des caracteres essentiels de l'animal, sa distinction du vegetal et des autres corps naturels, enfin, l'exposition des principes fondamentaux de la zoologie. Tome Deuxieme. Histoire des Polypes. Paris, J. B. Baillere, 684 pp.","Gordon, D. P., Taylor, P. D. & Bigey, F. P. (2009) Phylum Bryozoa. In: Gordon, D. P. (Ed.), New Zealand Inventory of Biodiversity. Volume One. Kingdom Animalia: Radiata, Lophotrochozoa, Deuterostomia. Christchurch, Canterbury University Press, pp. 271 - 297.","Busk, G. (1886) Report on the Polyzoa collected by H. M. S. Challenger during the years 1873 - 76. Part II. - The Cyclostomata, Ctenostomata, and Pedicellinea. Report on the Scientific Results of the Voyage of H. M. S. Challenger, Zoology, 17 [50], iviii, 1 - 47, 10 pls."]}

This paper describes 12 species of ctenostomes from the marine waters of the Korean Peninsula, including five new species—Alcyonidium bullitum n. sp., Alcyonidium pulposum n. sp., Alcyonidium busanensis n. sp., Immergentia cheongpodensis n. sp. and Penetrantia taeanata n. sp., the latter two constituting shell-borers that ramify within dead mollusk shells. Three previously described species are also newly added to the Korean fauna—Amathia acervata Lamouroux, 1824, Amathia medullaris Mawatari, 1972 and Walkeria prorepens Kubanin, 1992. The nomenclature of known species is updated and a revised checklist of all Korean Ctenostomata is included. New biological information is provided based on observation and microphotography of living and preserved colonies, SEM images of dry material and resin casts of ctenostome borings in mollusk shells.

Amathia medullaris Mawatari, 1972 (Fig. 18) Bowerbankia medullaris Mawatari, 1972: 300, figs 1, 2. Bowerbankia (Crassicaula) medullaris: Kubanin 1992: 26. Material examined. MBRBK1708, 36.6334�� N, 126.2997�� E, Cheongpodae, West Sea, 26 May 2017. MBRBK 1709, Daejin port, East Sea, 38.2957�� N, 128.2533�� E, 3 June 2017. Description. Colony encrusting brown alga Stephanocystis hakodatensis (Yendo) (formerly Cystophyllum hakodatense), on which it produces tufts of squat rounded sac-like zooids on stipes and fruiting bodies. Colony divided into two regions, an inner thick medullary region of branched stolons and single layer of almost spherical autozooids. At colony edges stolons thin and closely adherent to algal substratum, with erect autozooids sometimes biserially arranged along them (Fig. 18E). More-developed parts of colonies swell into thick complex mat of stolons and zooids, entire mat surrounding substratum but not tightly adherent to it (Fig. 18A). Zooidal orifice quadrate, polypide with distinct gizzard (note zooids in Fig. 18E). Zooids of Korean specimens with 10 evenly spaced tentacles all the same length (Fig. 18A, C, E). Eggs and embryos (Fig. 18B, D, G) yolky yellow color and brooded in zooids in which polypides had degenerated into brown bodies. Remarks. The colonies found by us match the original description of Mawatari (1972) except in two details. All the polypides in Korean colonies had 10 tentacles, whereas eight tentacles and orange eggs were described in the Japanese material. Since the eggs of our specimens were somewhat orange-yellow this detail may not matter, but tentacle number (eight or ten) is a significant species-specific character in all known Amathia species. Because species in this genus are so constrained in terms of tentacle number, if living Japanese specimens can be confirmed to have eight tentacles, the Korean material likely represents a new species. Amathia medullaris differs from all other known species in its production of a medullary region. Other 10-tentacled species, e.g. Amathia imbricata from the east and west Atlantic, may have thickly interwoven networks of stolons and zooids in well-grown colonies, but do not show the structural zonation found in A. medullaris. Kubanin (1992) created the subgenus Crassicaula for this species, based on the differences in its morphology in comparison with other species then classified in Bowerbankia. This distinction needs to be evaluated through further study. Distribution. Korea: Yellow Sea, East Sea. Also Peter the Great Bay, Russian coast of Japan Sea, and Hokkaido, Japan. Intertidal., Published as part of Seo, Ji-Eun, Chae, Hyun Sook, Winston, Judith E., Z��gor��ek, Kamil & Gordon, Dennis P., 2018, Korean ctenostome bryozoans-observations on living colonies, new records, five new species, and an updated checklist, pp. 251-283 in Zootaxa 4486 (3) on pages 275-279, DOI: 10.11646/zootaxa.4486.3.3, http://zenodo.org/record/1453233, {"references":["Mawatari, S. F. (1972) A new species of the genus Bowerbankia (Bryozoa, Ctenostomata) from Hokkaido. Journal of the Faculty of Science of Hokkaido University, Series 6, Zoology, 18, 300 - 304.","Kubanin, A. (1992) New bryozoan species (Gymnolaemata, Ctenostomata) from the Japan Sea. Zoologcheskii Zhurnal, 71, 19 - 31. [In Russian]"]}

Amathia distans Busk, 1886 (Figs. 22���23) Amathia distans Busk, 1886: 33, pl. 7, fig. 1; Vieira et al. 2008: 10; Fehlauer-Ale et al. 2011: 56, figs. 3, 4, 6, 8, 10 (cum syn.); Migotto et al. 2011: 269; Marques et al. 2013: 271; Vieira et al. 2014: 514; Almeida et al. 2015b: 3. Material examined. UFBA 1581, UFBA 2349���50, Todos os Santos Bay, 13��00���S, 38��32���W, 3���8 m, coll. 2013 (on sponge Callyspongia sp.); UFBA 1600, UFBA 2351, Camamu Bay, 13��53���S, 38��59���W, 18���20 m, coll. October 2012 (on sponge Dysidea etheria); UFBA 1617, UFBA 2352���53, Camamu Bay, 13��53���S, 38��59���W, 18���20 m, coll. October 2012 (on Mycale angulosa). Remarks. Fehlauer-Ale et al. (2011) redescribed A. distans and stated the bright yellow pigment spots in stolonal and zooidal surfaces, the thickly cuticularised slender stolon (Fig. 22), and the autozooids organized in clockwise and anticlockwise directions (Fig. 23) as distinctive characters of the species. Amathia distans has been reported on a variety of substrata such as algae, bryozoans and anthropogenic surfaces (Fehlauer-Ale et al. 2011). Here we present the first record of A. distans associated with the sponges Callyspongia sp., Dysidea etheria and Mycale angulosa. Distribution. Atlantic: Brazil (Alagoas, Bahia, Esp��rito Santo, Rio de Janeiro, S��o Paulo and Paran��) (Fehlauer-Ale et al. 2011)., Published as part of Almeida, Ana C. S., Souza, Facelucia B. C., Menegola, Carla & Vieira, Leandro M., 2017, Diversity of marine bryozoans inhabiting demosponges in northeastern Brazil, pp. 281-323 in Zootaxa 4290 (2) on pages 289-290, DOI: 10.11646/zootaxa.4290.2.3, http://zenodo.org/record/892719, {"references":["Busk, G. (1886) Report on the Polyzoa collected by H. M. S. Challenger during the years 1873 - 1876. Part II. The Cyclostomata, Ctenostomata and Pedicellinea. (17 th Report on the Scientific Results of H. M. S. Challenger during the years 1873 - 76, Zoology, 7 (50), i - viii + 1 - 47, 1 - 10 pls.","Vieira, L. M., Migotto, A. E. & Winston, J. E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810, 1 - 39.","Fehlauer-Ale, K. H., Vieira, L. M. & Winston, J. E. (2011) Molecular and morphological characterization of Amathia distans Busk and Amathia brasiliensis Busk (Bryozoa: Ctenostomata) from the tropical and subtropical Western Atlantic. Zootaxa, 2962, 49 - 62.","Migotto, A. E., Vieira, L. M. & Winston, J. E. (2011) Bryozoa. In: Amaral, A. C. Z. & Nallin, S. A. H. (Orgs.), Biodiversidade e ecossistemas bentonicos marinhos do Litoral Norte de Sao Paulo, Sudeste do Brasil. IB / UNICAMP, Campinas, pp. 265 - 272.","Marques, A. C., Kloh, A. S., Migotto, A. E., Cabral, A. C., Ravedutti, Tigo, A. P., Bettim, A. L., Razzolini, E. L., Cascon, J. M., Bardi, J., Kremer L. P., Vieira, L. M., Bezerra, L. E. A., Haddad, M. A., De Oliveira Filho, R. R., Gutierre, S. M. M., Miranda, T. P., Franklin, W. J. R. & Rocha, R. M. (2013) Rapid assessment survey for exotic benthic species in the Sao Sebastiao Channel, Brazil. Latin American Journal of Aquatic Research, 41, 265 - 285.","Almeida, A. C. S., Alves, O., Peso-Aguiar, M., Dominguez, J. & Souza, F. (2015 b) Gymnolaemata bryozoans of Bahia State, Brazil. Marine Biodiversity Records, 8, e 120."]}

As primary or obligate sessile organisms, bryozoans depend upon a substratum resource that affects their abundance, distribution and diversity. These animals can colonize virtually any type of substratum, including other organisms and artificial structures. Associations between bryozoans and sponges are commonly reported in the literature, but there are few studies discussing the association between these two taxa in detail. Here we present data on the bryozoan community found on shallow-water sponges from Bahia coast, northeastern Brazil, including their taxonomic status, colony form and adaptative structures utilized by these bryozoans to grow on sponges. Twenty-one bryozoan species were found attached to the surface of sixteen species of sponges. Five new species of cheilostome bryozoans are described. A total of 105 colonies were studied and most of them are erect delicate branching (44 colonies) and encrusting patches (34 colonies). The majority of bryozoan colonies were attached to the surface of rugose-textured sponges (67 colonies; 61%). This suggests that bryozoans are more likely to settle on irregular and rough surfaces. Patches colonies were mainly attached to the portion of the sponge that was in contact with the seabed, and spot colonies were particularly found in spatial refuges, showing the preference of larvae to settle on shaded and less exposed substrata. Most erect bryozoans were attached to the lateral sponge surface, other colonies grew on the underside and few on the upper surface of the sponges. These colonies were attached either using anchoring rhizoids, rigid bases, or stolons. The bryozoan species and genera reported here are common in northeastern Brazil and considered generalists in terms of larval settlement requirements. The bryozoan-sponge association studied is considered a non-obligatory commensalism (inquilinism).

Zoobotryon verticillatum, a widely dispersed stoloniferous fouling bryozoan, has been recently recorded from specimens collected from the hulls of vessels berthed in an Israeli marina. Though conspicuous and distinctive, it has been rarely recorded in surveys of the bryozoan fauna along the Levant coastline conducted over the past half century. It is suggested the species is native to the Caribbean Sea, and an invasive alien elsewhere, including the Mediterranean Sea.

Morphological and molecular analyses have proven to be complementary tools of taxonomic information for the redescription of the ctenostome bryozoans Amathia brasiliensis Busk, 1886 and Amathia distans Busk, 1886. The two species, originally described from material collected by the ‘Challenger’ expedition but synonymized by later authors, now have their status fixed by means of the selection of lectotypes, morphological observations and analyses of DNA sequences described here. The morphological characters allowing the identification of living and/or preserved specimens are (1) A. brasiliensis: whitish-pale pigment spots in the frontal surface of stolons and zooids, and a wide stolon with biserial zooid clusters growing in clockwise and anti-clockwise spirals along it, the spirality direction being maintained from maternal to daughter stolons; and (2) A. distans: bright yellow pigment spots in stolonal and zooidal surfaces including lophophores, and a slender stolon, thickly cuticularized, with biserial zooid clusters growing in clockwise and anti-clockwise spirals along it and the spirality direction not maintained from maternal to daughter stolons. Pairwise comparisons of DNA sequences of the mitochondrial genes cytochrome c oxidase subunit I and large ribosomal RNA subunit revealed deep genetic divergence between A. brasiliensis and A. distans. Finally, analyses of those sequences within a Bayesian phylogenetic context recovered their genealogical species status.

Amathia (Amathia) pulchra Miers 1885: 589. Anamathia pulchra Miers 1886: 26, pl. IV, fig. 1. Remarks. Originally assigned to Amathia (Miers 1885), but ultimately to ��� Anamathia S.I. Smith, 1884 ��� (Miers 1886), Scyramathia A. Milne-Edwards, 1881 (Alcock 1895) and Rochinia A. Milne-Edwards, 1875 (Sakai 1938). None of these revisions cite Miers 1885. Alphonse Milne-Edwards & Bouvier (1900: 131) cited ��� 1881. Scyramathia. A. Milne-Edwards, C. R. Acad. des Se., 5 d��c. 1881 ��� and this is followed by Alcock (1895: 201). Rathbun (1925: 204) cited 1880 as the authority date for Scyramathia and this appears to be correct as the BL copy (call number AC. 424) of Comptes Rendus Hebdomadaires des S��ances de l'Acad��mie des Sciences Volume 91, sheets 33���58, pages 251���448, containing the paper of A. Milne-Edwards (1880 b) is dated with a red British Museum Stamp ��� 6 OC 80 ���. This availability date in London confirms Scyramathia A. Milne-Edwards, 1880. Furthermore, volume 7 of the Proceedings of the U.S. National Museum has the replacement name Anamathia S.I. Smith for the preoccupied Amathia Roux, 1828 and was actually published in 1885 (ICZN Opinion 712). Valid as Rochinia pulchra (Miers, 1885)., Published as part of Froglia, Carlo & Clark, Paul F., 2011, The forgotten Narrative of H. M. S. Challenger and the implications for decapod nomenclature, pp. 45-56 in Zootaxa 2788 on page 51, DOI: 10.5281/zenodo.200635, {"references":["Alcock, A. (1895) Materials for a carcinological fauna of India, 1. The Brachyura Oxyrhyncha. Journal of the Asiatic Society of Bengal, (Part 2, Natural Sciences), 64, 157 - 291, pls 3 - 5.","Milne-Edwards, A. (1875) Etudes sur les xiphosures et les crustaces podophthalmaires. In: Mission scientifique au Mexique et dans l'Amerique centrale, ouvrage publie par ordre du Ministre de l'Instruction publique. Recherches zoologiques pour servir a l'histoire de la faune de l'Amerique centrale et du Mexique, publiees sous la direction de M. H. Milne Edwards, membre de l'Institut. Cinquieme partie. Paris, Imprimerie nationale. Livraison 3, 57 - 120, pls 15 - 20. [for dates see Crosnier and Clark, 1998].","Sakai, T. (1938) Studies on the crabs of Japan III. Brachygnatha, Oxyrhyncha. Tokyo, Yokendo, pp. 194 - 364, pls XX-XLI.","Milne-Edwards, A. & Bouvier, E. L. (1900) Crustaces Decapodes. Premiere partie. Brachyures et Anomoures. Expeditions Scientifiques du Travailleur et du Talisman pendant les annees 1880, 1881, 1882, 1883. Ouvrage publie sous les auspices du ministere de l'instruction publique sous la direction de A. Milne-Edwards membre de l'Institut, president de la Commission des Dragages sous-marins, Directeur du Museum d'Histoire Naturelle. Libraires de l'Academie de Medecine, Paris, 1, 1 - 396, pls I-XXXII.","Rathbun, M. J. (1925) The spider crabs of America. Bulletin of the United States National Museum, 129, xx + 1 - 611, pls 1 - 283.","Milne-Edwards, A. (1880 b) Compte rendu sommaire d'une exploration zoologique faite dans le golfe de Gascogne, a bord du navire de l'Etat le Travailleur. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, Paris, 91, 355 - 360.","Roux, P. (1828) Crustaces de la Mediterranee et de son littoral. Pages not numbered; parts 1, 2, pls 1 - 10. Marseille, Imprimerie D'Achard. [for dates see Monod, 1956]"]}

Amathia distans Busk, 1886 (Figs 3, 4, 6, 8, 10; Tables 1 and 3) Amathia distans Busk, 1886: 33, pl. 7, fig. 1; Marcus 1937: 134 (part) (not pl. 27, fig. 72); 1941: 28, fig. 29; 1949: 27, fig. 42; 1955: 313 (part); Osburn 1940: 339; Winston 1982: 110, fig. 10; d���Hondt 1983: p. 65 (part; form 1, p. 69). Material examined. Lectotype (chosen here): NHMUK 1887.12. 9.925 (Figures 3 and 4), Amathia distans (G. Busk det.), H.M.S. ��� Challenger ���, Bahia (Brazil), 10���20 fms (18.29���36.58 m), figured by Busk (1886, pl. vii, fig. 1). Paralectotypes: NHMUK 1887.12. 9.926, Amathia distans (G. Busk det.), H.M.S. ��� Challenger ���, Bahia (Brazil), 10���20 fms (18.29���36.58 m); NHMUK 1899.7.1.4358��� 4359, Amathia distans, H.M.S. ��� Challenger ���, Bahia (Brazil), G. Busk Collection; NHMUK 1963.2. 12.359, Amathia distans, H.M.S. ��� Challenger ���, Bahia (Brazil), Dundee Collection. Additional material (Table 1): MZUSP (498���500), MZUSP (508���510) (donated to NHMUK), S��o Sebasti��o, S��o Paulo State, Brazil. Redescription. Colonies cuticularized, erect, regularly branched, transparent white to yellowish tan. Frontal surface of stolons and zooids of living colonies with bright yellow pigment spots. Autozooids in biserial clusters, 9 to 19 pairs of long clockwise and/or anticlockwise complete spirals at the distal end around the thickly cuticularized slender stolon; direction of spirals not maintained from maternal to daughter stolons; each maternal stolon buds two daughter stolons, one of which has autozooidal clusters organized in a clockwise spiral and the other in an anticlockwise spiral. Clusters present from distal half of stolon to three quarters of its length. Zooids tubular, connate for almost their entire length, not varying in size with the position of the series. Orifice terminal, circular. Polypides with 8 equal, slightly curved tentacles about 0.23 mm in length, with yellow pigment spots. Remarks. Amathia distans is distinguished from its congeners by the combined presence of bright yellow pigment spots in stolonal and zooidal surfaces, including lophophores, and by a thickly cuticularized slender stolon (0.09���0.14 mm in diameter). The daughter stolons have autozooids organized in clockwise and anticlockwise directions, with the direction of spirals rarely maintained from maternal to daughter stolons as in A. brasiliensis. Amathia distans and A. brasiliensis, which were described for the same locality by Busk (1886), were originally distinguished by the colony branching pattern, the stolon diameter and the size and position of the zooid clusters on the stolon. These distinctive characteristics have been confirmed through morphological analyses of syntype specimens deposited at the NHMUK. Here, we have selected and figured (Figs 1���4) the lectotype specimens of both A. brasiliensis (NHMUK 1887.12.9.928) and A. distans (NHMUK 1887.12.9.925). Specimens attributed to A. distans by different researchers over the years show morphological characteristics that do not completely fit the original description of Busk (1886) and probably pertain to other species. Unfortunately, any potential pigmentation differences cannot be observed in preserved colonies, as the pigment is lost in alcohol; only the characters of zooids, zooid clusters and internodes are available for comparison. Harmer (1915) identified as A. distans specimens characterized by dichotomous colonies with zooids partially joined, usually forming one complete loop in the distal part of the slender internode, about 0.10 mm in diameter. Although the diameter of the stolons measured by Harmer (1915) is similar to that in A. distans Busk, the Siboga specimens have zooids unjoined for two-fifths or more of their length. Several Brazilian colonies with both wider and slenderer stolons were identified by Marcus (1937, 1941, 1949, 1955) under the name A. distans. Figures based upon specimens collected in S��o Paulo State (Marcus 1937: pl. 27, fig. 72; Marcus 1949: fig. 42) show stolons 0.15���0.35 mm in diameter and were probably based upon both A. distans and A. brasiliensis. Osburn (1940) distinguished Caribbean specimens of A. brasiliensis and A. distans on the basis of stolon diameter of stolon and the degree of zooid joining. Yellow-pigmented zooids were first noted in A. distans collected in Florida by Winston (1982), and have never been reported in specimens of A. distans from the Pacific (D.P. Gordon, pers. comm.). The yellow-spotted Atlantic variety of A. distans (footnote 4, p. 69 in d���Hondt 1983) is considered A. distans sensu stricto. D���Hondt���s figured specimen (d���Hondt 1983: p. 64, fig. 35 C; p. 69, form 2) probably represents a different species, based on the presence of autozooids 0.20���0.25 mm in diameter, wider than those of A. distans Busk (Table 3). The small specimen from South Australia recorded as A. distans by MacGillivray (1895) has unjoined zooids; this specimen also probably belongs to a different species, which resembles ��� Amathia distans form 2 ��� of d���Hondt (1983: p. 69). Chimonides (1987) discussed other morphological similarities (e.g. shape of clusters of zooids rotating 360 �� around stolon) and differences (proportion of bare stolon between adjacent zooid clusters and diameter of stolon) between A. brasiliensis and A. distans. Souto et al. (2010) reported a similar appearance amongst A. minoricensis and some widespread species, including A. distans. However, according to those authors, A. minoricensis is readily distinguished from either A. distans or A. brasiliensis by the presence of rhizoids, autozooidal shape, degree of twisting of zooid clusters along stolons and the branching pattern of colonies. Biological notes. Amathia distans sensu stricto occurs at least from Brazil to Florida on substrata including algae, bryozoans, and anthropogenic surfaces. NHMUK MZUSP MZUSP MZUSP MZUSP MZUSP 1887.12. 9.925 498 499 508 509 510 Autozooid length Min-Max - 0.310���0.540 0.350���0.470 0.330���0.480 0.390���0.520 0.310���0.520 Mean (SD) - 0.450 (0.070) 0.405 (0.032) 0.414 (0.041) 0.424 (0.039) 0.410 (0.060) Autozooid width Distribution. The species was originally described from the Brazilian State of Bahia by Busk (1886: p. 33), and was reported for the shores of the States of Esp��rito Santo, Rio de Janeiro, S��o Paulo, Paran�� (Marcus 1937, 1941, 1949, 1955), and Alagoas (Vieira et al. 2007, 2008). It has since been treated as a widespread fouling species in warm waters, with records for Australia, Indonesia, Florida, Southern California, Gulf of California, and New Zealand (viz. Harmer 1915; Gordon & Mawatari 1992; Soule in Osburn 1953; Menon 1972; Gordon, 2009; K. Tilbrook pers. comm. 2010.). However, some of these records are doubtful and require reinvestigation., Published as part of Fehlauer-Ale, Karin H., Vieira, Leandro M. & Winston, Judith E., 2011, Molecular and morphological characterization of Amathia distans Busk and Amathia brasiliensis Busk (Bryozoa: Ctenostomata) from the tropical and subtropical Western Atlantic, pp. 49-62 in Zootaxa 2962 on pages 56-58, DOI: 10.5281/zenodo.208030, {"references":["Busk, G. (1886) Report on the Polyzoa collected by H. M. S. Challenger during the years 1873 - 1876. Part. 2. The Cyclostomata, Ctenostomata and Pedicellinea. Report on the Scientific Results of the Voyage of the H. M. S. \" Challenger \", Zoology, 17 (3), 1 - 47.","Marcus, E. (1937) Bryozoarios marinhos brasileiros I. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Zoologia, 1, 5 - 224.","Osburn, R. C. (1940) Bryozoa of Porto Rico with a resume of the West Indian Bryozoan fauna. Scientific Survey of Porto Rico and the Virgin Islands, 16, 321 - 486.","Winston, J. E. (1982) Marine bryozoans (Ectoprocta) of the Indian River area, Florida. Bulletin of the American Museum of Natural History, 173, 99 - 176.","d'Hondt, J. - L. (1983) Tabular keys for the identification of the Recent ctenostomatous Bryozoa. Memoires de l'Institut Oceanographique, 14, 1 - 134.","Harmer, S. F. (1915) The Polyzoa of the Siboga Expedition. Part 1. Entoprocta, Ctenostomata and Cyclostomata. Siboga Expedition Reports, 28 A, 1 - 180.","Marcus, E. (1941) Briozoarios marinhos do litoral paranaense. Arquivos do Museu Paranaense, 1, 7 - 36.","Marcus, E. (1949) Some Bryozoa from the Brazilian coast. Comunicationes Zoologicas del Museo de Historia Natural de Montevideo, 3 (53), 1 - 33.","Marcus, E. (1955) Notas sobre briozoos marinhos brasileiros. Arquivos do Museu Nacional do Rio de Janeiro, 42, 273 - 341.","MacGillivray, P. H. (1895) On the Australian species of Amathia. Proceedings of the Royal Society of Victoria, n. s., 7, 131 - 140.","Chimonides, P. J. (1987) Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). Bulletin of the British Museum (Natural History), Zoology, 52, 307 - 358.","Souto, J., Fernandez-Pulpeiro, E. & Reverter-Gil, O. (2010) The genus Amathia Lamouroux (Bryozoa: Ctenostomata) in Iberian waters. Cahiers de Biologie Marine, 51, 181 - 195.","Vieira, L. M., Gordon, D. P. & Correia, M. D. (2007) First record of a living ditaxiporine catenicellid in the Atlantic, with a description of Vasignyella ovicellata n. sp. (Bryozoa). Zootaxa, 1582, 49 - 58.","Vieira, L. M., Migotto, A. E. & Winston, J. E. (2008) Synopsis and annotated checklist of recent marine Bryozoa from Brazil. Zootaxa, 1810, 1 - 39.","Gordon, D. P. & Mawatari, S. F. (1992) Atlas of marine-fouling Bryozoa of New Zealand ports and harbours. Miscellaneous Publications of the New Zealand Oceanographic Institute, 107, 1 - 52.","Osburn, R. C. (1953) Bryozoa of the Pacific coast of America, part. 3, Cyclostomata, Ctenostomata, Entoprocta and Addenda. Allan Hancock Pacific Expeditions, 14, 613 - 841.","Menon, N. R. (1972) Species of the sub-order Ctenostomata Busk (Bryozoa) from Indian waters. Internationale Revue der gesamten Hydrobiologie und Hydrographie, 57, 599 - 629.","Gordon, D. P. (2009) Baudina gen. nov., constituting the first record of Pasytheidae from Australia, and Sinoflustridae fam. nov., with a checklist of Bryozoa and Pterobranchia from Beagle Gulf. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 25, 43 - 54."]}

The Narrative of the Challenger cruise was published in 1885 and contained valid descriptions of crustacean decapod taxa. While C.S. Bate (Challenger Macrura) mentioned the discovery of new taxa without making them valid, J.R. Henderson (Anomura) and E.J. Miers (Brachyura) provided short descriptions including figures that made several new names available. With a few exceptions, these valid scientific names together with their date of availability have been overlooked by almost all subsequent taxonomic studies. Consequently the decapod taxa named in the Challenger Narrative are listed and the implications for nomenclature in each case are discussed.

Amathia brasiliensis Busk, 1886 (Figs 1, 2, 5, 7, 9; Tables 1���2) Amathia brasiliensis Busk, 1886: 34, pl. 7, fig. 2; Osburn 1940: 339. Amathia distans: Marcus 1937: 134 (part), pl. 27, fig. 72; 1955: 313 (part); Maturo 1957: 23, fig. 12; d���Hondt 1983: p. 65 (part). Non Busk 1886: 33. Material examined. Lectotype (chosen here): NHMUK 1887.12. 9.928 (slide mounted by A.B. Hastings using Borax-carmine Glycerine; Figures 1 and 2), Amathia brasiliensis (G. Busk det.), H.M.S. ��� Challenger ���, Bahia (Brazil), 10���20 fms (18.29���36.58 m). Paralectotypes: NHMUK 1887.12. 9.927, Amathia brasiliensis (G. Busk det.), H.M.S. ��� Challenger ���, Bahia (Brazil), 10���20 fms (18.29���36.58 m); NHMUK 1890.4. 14.5, Amathia brasiliensis, Bahia (Brazil), G. Busk Supplementary Collection; NHMUK 1899.7. 1.4520, Amathia brasiliensis, H.M.S. ��� Challenger ���, Bahia (Brazil), G. Busk Collection; NHMUK 1963.2. 12.356, Amathia brasiliensis, H.M.S. ��� Challenger ���, Bahia (Brazil), Dundee Collection. Additional material (Table 1): MZUSP 495, MZUSP 505 (donated to NHMUK), Macei��, Alagoas State, Brazil; MZUSP (492, 493), MZUSP (504, 506) (donated to NHMUK), S��o Sebasti��o, S��o Paulo State, Brazil; MZUSP 494, Ilhabela, S��o Paulo; MZUSP (496, 497), MZUSP 507 (donated to NHMUK), S��o Francisco do Sul, Santa Catarina State, Brazil. Redescription. Colonies cuticularized, erect, densely branched, brownish. Frontal surface of stolons and zooids delicate, with inconspicuous whitish-pale pigment spots observed only in living colonies; colonies in alcohol whitish-brown in color. Autozooids in biserial clusters, 8 to 18 pairs in long complete clockwise or anticlockwise spirals along a wide stolon; the direction of spirals maintained from maternal to daughter stolons. Clumps occupying from half to three quarters of distal portion of stolon. Autozooids tubular, most of them almost completely unjoined, rarely joined at one third of their total length, not varying in size. Orifice terminal, approximately subquadrangular. Rhizoids sometimes present, arising in the proximal end of stolon. Polypides with 8 equal straight tentacles, measuring approximately 0.27 mm in length. Remarks. Amathia brasiliensis was originally described from Bahia State, Brazil (Busk 1886). This species is characterized by whitish-pale pigment spots in the frontal surface of stolons and zooids, and by a wide stolon (about 0.22 mm of diameter) with biserial zooid clusters growing in clockwise or anticlockwise spirals along it. Busk (1886) noted a tendency of direction of spirals, characteristic of this species. Osburn (1940) disagreed with the synonymization of the name A. brasiliensis under A. distans adopted by Marcus (1937), and used the epithet brasiliensis for specimens with wider stolons (0.18���0.30 mm). The North Carolina specimens reported by Maturo (1957) under the name A. distans have a wider stolon than A. distans from Brazil (see Tables 2���3 for comparison), and hence probably belong to A. brasiliensis. Specimens from the Suez Canal collected by Hastings (1927), registered as Amathia ? brasiliensis (NHMUK 1926.9.6.25), have stems with similar diameters (0.18���0.20 mm) to those of A. brasiliensis Busk, but have smaller and broader free ends of zooids than the ���Challenger��� specimens of A. brasiliensis. D���Hondt (1983) redescribed the Suez Canal material as Amathia distans var. aegyptiana, which was later raised to specific rank as Amathia aegyptiana by Chimonides (1987). The rooting processes are present in both A. brasiliensis and A. aegyptiana and the long bare terminal ramification, rarely present in A. brasiliensis, was not described for A. aegyptiana. Chimonides (1987), although noticing some similarities among Amathia pruvoti, A. distans, A. aegyptiana and A. brasiliensis, observed several morphological characteristics that support each one as a distinct entity. These include the spiraling pattern of autozooidal groups (close to 360 �� in the last three species), the production of rhizoids only by A. brasiliensis, and the autozooidal spirality direction between maternal and daughter stolons. According to Chimonides (1987), A. aegyptiana is the only one that produces autozooidal groups in which the direction of the spirality is maintained. Another distinctive character of A. aegyptiana is the absence of rhizoids [according to Chimonides (1987) and Souto et al. (2010)], but this structure is inconstant among the different specimens of A. brasiliensis studied in the Western Atlantic and could represent ecophenotypical variation. Autozooid length Min-Max - 0.320���0.580 0.300��� 0.680 0.340���0.540 0.390���0.520 0.410���0.570 Mean (SD) - 0.422 (0.068) 0.300 (0.124) 0.450 (0.058) 0.440 (0.046) 0.454 (0.050) Autozooid width Souto et al. (2010) described Amathia minoricensis from Balearic waters (Iberian Peninsula), in which the direction of twisting remains the same as that of the maternal stolons. Although A. brasiliensis and A. minoricensis present overlapping measurements for autozooids and stolon width [page 189 of Souto et al. (2010); Table 2 of this study], the latter species is readily distinguished by the helix of autozooids describing about 180���250 ��, the autozooids being joined for almost their total length, and the length of the stolon and zooid cluster being shorter in A. brasiliensis than in A. minoricensis. Souto et al. (2010) also noted differences between the spiraling direction of A. aegyptiana and A. brasiliensis, which according to them is not conserved along the colony of the latter species. However, in all specimens of A. brasiliensis here analyzed, including the type specimens, the direction of twisting is conserved along the colony. Busk (1886: p. 34) also noted for A. brasiliensis that the ���comparative distinctness of the zooecia in the spiral series shows a tendency in the same direction���. Biological notes. In Brazil, A. brasiliensis is common on various substrata, including algae, bryozoans and anthropogenic surfaces, and was observed also on or with other vesiculariids, such as A. distans, Amathia cf. vidovici, Amathia cf. crispa and Z. verticillatum. Distribution. Western Atlantic: North Carolina (Maturo 1957), Bermuda, Porto Rico (Osburn 1940), and Brazil (Vieira et al. 2008). Brazil: Fernando de Noronha Island, and states of Alagoas, Bahia, S��o Paulo and Santa Catarina., Published as part of Fehlauer-Ale, Karin H., Vieira, Leandro M. & Winston, Judith E., 2011, Molecular and morphological characterization of Amathia distans Busk and Amathia brasiliensis Busk (Bryozoa: Ctenostomata) from the tropical and subtropical Western Atlantic, pp. 49-62 in Zootaxa 2962 on pages 52-56, DOI: 10.5281/zenodo.208030, {"references":["Busk, G. (1886) Report on the Polyzoa collected by H. M. S. Challenger during the years 1873 - 1876. Part. 2. The Cyclostomata, Ctenostomata and Pedicellinea. Report on the Scientific Results of the Voyage of the H. M. S. \" Challenger \", Zoology, 17 (3), 1 - 47.","Osburn, R. C. (1940) Bryozoa of Porto Rico with a resume of the West Indian Bryozoan fauna. Scientific Survey of Porto Rico and the Virgin Islands, 16, 321 - 486.","Marcus, E. (1937) Bryozoarios marinhos brasileiros I. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Zoologia, 1, 5 - 224.","Maturo, F. J. S. (1957) Study of the Bryozoa of Beaufort, North Carolina, and vicinity. Journal of the Elisha Mitchell Scientific Society, 73, 11 - 68.","d'Hondt, J. - L. (1983) Tabular keys for the identification of the Recent ctenostomatous Bryozoa. Memoires de l'Institut Oceanographique, 14, 1 - 134.","Hastings, A. B. (1927) Zoological results of the Cambridge expedition to the Suez Canal, 1924, 20. Report on the Polyzoa. Transactions of the Zoological Society of London, 22, 331 - 353.","Chimonides, P. J. (1987) Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). Bulletin of the British Museum (Natural History), Zoology, 52, 307 - 358.","Souto, J., Fernandez-Pulpeiro, E. & Reverter-Gil, O. (2010) The genus Amathia Lamouroux (Bryozoa: Ctenostomata) in Iberian waters. Cahiers de Biologie Marine, 51, 181 - 195.","Vieira, L. M., Migotto, A. E. & Winston, J. E. (2008) Synopsis and annotated checklist of recent marine Bryozoa from Brazil. Zootaxa, 1810, 1 - 39."]}

Souto, J., Fernández-Pulpeiro, E., Reverter-Gil, O. (2011): Presence of rhizoids in two species of the genus Bowerbankia (Bryozoa: Ctenostomata) and their systematic relevance. Journal of Natural History 45 (41-42): 2543-2557, DOI: 10.1080/00222933.2011.597003, URL: http://dx.doi.org/10.1080/00222933.2011.597003