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The accumulation of plastic litter in coastal environments has become an issue of high priority for policymakers around the globe, due to their potential hazardous effects to biota and human health, and their impact on ecosystem services and local economies. To develop effective mitigation measures, it is critical to acquire knowledge on the distribution and levels of plastic litter. However, in many world regions, such as in West Africa or Southeast Asia, the concentration of plastics reaching coastal areas is still poorly documented. To address the data gaps in marine plastic litter distribution worldwide, citizen science programs are instrumental in complementing shoreline assessments, and are effective in increasing public awareness of plastic pollution. The Citizen Observation of Local Litter in coastal ECosysTems (COLLECT) project is a citizen science initiative which aims to acquire distribution and abundance data of coastal plastic debris in seven countries, in Africa (Benin, Cabo Verde, Côte d'Ivoire, Ghana, Morocco, Nigeria) and Asia (Malaysia). The project consists of training local students (15-18 years old) from secondary cycle institutions on sampling and analysing macro-, meso- and microplastic in beach sediments, using scientific procedures. The project will also measure the impact of the citizen science intervention by assessing shifts in ocean literacy and pro-environmental behaviour in the students. The results from COLLECT will contribute to establishing baseline information on coastal plastic debris, with citizen science being an enabler of open science, allowing data to be freely available to the public, academics and policymakers. Results will further contribute to the identification of hotspots of plastic coastal litter, and bring awareness to local communities on the potential consequences of plastic pollution. Also see: https://micro2022.sciencesconf.org/427338/document, In MICRO 2022, Online Atlas Edition: Plastic Pollution from MACRO to nano

Outcome from working on the United Nations Environment Programme Advisory Group with the aim to address the UN Environment Assembly’s adopted resolution (UN/EA.4/RES.6) on Marine Plastic Litter and Microplastics by recommending indicators to harmonise monitoring and assessment and informing on policies and action environmentally sound technology innovations.

Diversity of the family Fungiidae was investigated in Malaysian waters during an expedition in 2004. From the samples collected on 18 reef locations around Malaysian waters, only 12 locations contained fungiid corals. From the locations that have fungiid corals, 11 species from five genera were recorded. They are Fungia (Ctenactis) echinata, Fungia (Ctenactis) simplex, Fungia (Danafungia scruposa, Fungia (Danafungia) danai, Fungia (Fungia) fungites, Fungia (Pleuractis) paumotensis, Fungia (Pleuractis) scutaria, Halomitra pileus, Sandalolitha robusta, Herpolitha limax and Polyphyllia talpina. The diversity of the fungiid corals at Hempasan Dang Ajar and Terumbu Peninjau far outweighs all other sites with six different species at each location. Other locations have three species on average. Out of 11 species of Fungiidae coral, Sandalolitha robusta and Herpolitha limax are the most commonly found species. They are found on seven sampling locations each. The rarest species are Fungia simplex, Fungia scruposa and Fungia paumotensis which can be found at only one sampling location each. Other available species could be found at three locations on average. This study has reported higher number of fungiid species compared to previous studies done in Peninsular Malaysia but similar with the study done in Singapore.

Acanthodesia cf. irregulata (Liu, 1992) comb. nov. Fig. 3 G���L cf. Membranipora irregulata Liu, 1992: 124, figs 14���17. Biflustra irregulata ��� Gordon et al. 2007: 46, fig. 1d. Material MALAYSIA: MSL BRY005, Pulau Betong, Penang, on oyster rafts. MSL BRY006, Balik Pulau, Penang, on plastic. MSL BRY007, BRY026, Batu Maung, Penang, on gastropod shells found among fishing debris. Description Colony encrusting, multiserial, unilamellar. Ancestrula twinned, proximally overgrown by later zooids in the example scanned. Autozooids subhexagonal, gently convex distally, 0.36���0.54 mm long by 0.22���0.30 mm wide; zooidal boundaries marked by a fine fissure; mural rim salient, narrow to moderately wide, pustulose; opesia occupying nearly all frontal surface, ovoidal, sometimes pear-shaped; gymnocyst absent or represented by low, rounded tubercles at proximal and proximolateral corners of zooids (Fig. 3 K���L); cryptocyst narrow, broadest proximally where it forms a shallow planar shelf, variable in width laterally, pustulose. Kenozooids developed at growth irregularities, diamond-shaped, variable in size, opesia teardrop-shaped. Remarks Acanthodesia irregulata was originally described from the South China Sea and subsequently identified by Gordon et al. (2007) from Bangladesh. Two characteristic features were noted by Liu (1992): the presence of cuticular spinules on the frontal membrane and operculum, and tubercles at the proximal corners of some of the zooids. The former have not been observed in the studied material from Malaysia, which is bleached. However, tubercles do occur in some of the studied Malaysian specimens (Fig. 3 K���L) and the size of the autozooids is very similar to that given by Gordon et al. (2007) in their Bangladeshi material. The paucity of taxonomic characters in the skeleton of the Malaysian specimens precludes certain identification as A. irregulata. It should also be noted that more than one species may be represented in Penang to judge from the variations in skeletal morphology; for example, tubercles can be present (Fig. 3 K���L) or absent (Fig. 3 G���J), and the width of the mural rim varies between specimens (Fig. 3 G���L)., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on page 9, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Liu X. 1992. On the genus Membranipora (Anasca: Cheilostomata: Bryozoa) from south Chinese seas. Raffles Bulletin of Zoology 40: 103 - 144.","Gordon D. P., Maruf Hossain M. M. & Wood T. S. 2007. The known and anticipated bryozoan diversity of Bangladesh. Journal of Taxonomy and Biodiversity Research 1 (2): 45 - 58."]}

Celleporaria aperta (Hincks, 1882) Fig. 10 A���F Schizoporella aperta Hincks, 1882: 126, pl. 5, fig. 3. Celleporaria aperta ��� Gordon et al. 2007: 49, fig. 2f���g. Material MALAYSIA: MSL BRY017a, Kuah jetty, Langkawi, fouling mussel shell attached to rope hanging from jetty. Description Colony encrusting, multiserial, becoming multilamellar through frontal budding. Basal autozooids (Fig. 10B) subhexagonal, 0.38���0.53 mm long by 0.20���0.28 mm wide; frontally budded autozooids (Fig. 10C) irregularly shaped, about 0.34 mm long by 0.28 mm wide; zooidal boundaries marked by fine fissures; frontal shield gently convex, smooth when first formed, becoming pustulose, peripheral ring of large subcircular areolar pores; orifice generally inclined distally, about 0.15 mm long by 0.13 mm wide, with a distinct rounded sinus (Fig. 10D), a pair of condyles present in proximal part of anter; two short, widely separated oral spines with closed ends can occur (Fig. 10D); ovicell hood-like (Fig. 10E), broad, about 0.10 mm long by 0.20 mm wide, surface with pustules, non-porous.Adventitious avicularia located suborally (Fig. 10 D���E), slightly eccentric to median axis, frontal plane almost perpendicular to surface of autozooid, small; rostrum rounded, tip dentate; cross-bar calcified. Interzooidal avicularia (Fig. 10F) distributed irregularly among frontally budded autozooids, variable in size, up to 0.32 mm long by 0.15 mm wide, distal end slightly raised; rostrum spatulate, rounded, distally dentate; cross-bar calcified, some with small lingulum; opesia semielliptical, wider than long. Remarks Malaysian specimens of Celleporaria aperta clearly show the dentate distal tips of the avicularian rostra (Fig. 10F), like those depicted by Hincks (1882: pl. 5, fig. 3), but not apparent in all material attributed to the species by subsequent authors. Hincks (1882: 126) gave the provenance of his material as ��� Singapore or Philippines ��� and the species is regarded as widespread in the tropical and subtropical Indo-Pacific (Gordon et al. 2007)., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 18-19, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Hincks T. 1882. Contributions towards a general history of the marine Polyzoa. IX. Foreign Cheilostomata (Miscellaneous). Annals and Magazine of Natural History, Series 5, 9: 116 - 127. http: // dx. doi. org / 10.1080 / 00222938209459003","Gordon D. P., Maruf Hossain M. M. & Wood T. S. 2007. The known and anticipated bryozoan diversity of Bangladesh. Journal of Taxonomy and Biodiversity Research 1 (2): 45 - 58."]}

Parasmittina winstonae Liu et al., 2001 Fig. 11 A���L Parasmittina winstonae Liu et al., 2001: 623, pl. 55, figs 1���7. Parasmittina winstonae ��� Tilbrook 2006: 156, pl. 29b, 30d���e. Material MALAYSIA: MSL BRY018, Kuah jetty, Langkawi, several colonies encrusting bivalves growing on a rope hanging from jetty. MSL BRY019a, Kuah jetty, Langkawi, encrusting barnacles and bivalve growing on a rope hanging from jetty. MSL BRY020a, BRY021a, Pulau Betong, Penang, encrusting barnacles from oyster rafts. Description Colony encrusting, multiserial, unilamellar; growing edge developing giant buds (Fig. 11J). Ancestrula tatiform (Fig. 11C), about 0.22 mm long by 0.17 mm wide, with subcircular opesia 0.16 mm long by 0.14 mm wide surrounded by 9 spines, becoming overgrown in older colonies; periancestrular zooids numbering 5 or 6, 0.26���0.32 mm long by 0.22���0.30 mm wide, some bearing adventitious avicularia, calcification of their frontal shields extending as short lobes between oral spines of ancestrula (Fig. 11D). Autozooids subhexagonal, distally rounded, 0.32���0.45 mm long by 0.22���0.33 mm wide; boundary walls salient with a fine fissure; frontal shield gently convex, coarsely pustulose, about 15���20 marginal areolar pores separated by ridges, areolae impinging on central part of shield in older zooids (Fig. 11H), no pseudopores; primary orifice egg-shaped to rounded rectangular, 0.10���0.11 mm long by 0.10 mm wide, large relative to zooid size, lappets developed distolaterally, lyrula narrow, 17���20 ��m high by 25���37 ��m wide at top edge, condyles present about one-third distance along orifice, 15 ��m high, medioproximal edge serrated (Fig. 11K); oral spine bases usually numbering two, occasionally one or lacking; ovicell hyperstomial (Fig. 11 H���I), broader than long, 0.13���0.16 mm long by 0.18 mm wide, containing 25���40 pores with raised rims, variable in size, circular, elliptical or dumbbell-shaped, margins overgrown by cryptocystal calcification. Avicularia adventitious, of three types: small acuminate, small plectrum-shaped, and giant. Small acuminate avicularia variously located (Fig. 11G, I), increasing in number during ontogeny, usually directed distally or distolaterally to laterally and towards midline of zooid, occasionally proximolaterally, about 0.09���0.12 mm long by 0.05���0.06 mm wide; rostrum acuminate, a high triangle, shallow rostral shelf; cross-bar calcified, straight or slightly concave; opesia semicircular. Small plectrum-shaped avicularia located laterally or distolaterally of orifice (Fig. 11K), occasionally in more proximal positions around perimeter of autozooid, directed and inclined outwardly, narrow proximally, broadening distally, about 0.05���0.06 mm long by 0.05���0.08 mm wide (measured in plane of colony surface); rostrum rounded trapezoidal, shallow rostral shelf; cross-bar completely or incompletely calcified, straight; opesia semicircular, small. Giant avicularia occasionally present (Fig. 11F), directed distally, originating proximolaterally and passing laterally of orifice, which may be torqued, extending over frontal shield of distal zooid, straight or curved towards autozooid midline, 0.35���0.40 mm long by 0.11���0.13 mm wide; rostrum spatulate, long, with a deep shelf occupying onethird to two-thirds of length; cross-bar slightly concave; opesia semicircular. Remarks Parasmittina is a very diverse genus with a global distribution. Different species are best characterised by their avicularia, which are often of more than one morph within a single colony. First described formally from South China (Liu et al. 2001), Parasmittina winstonae has three kinds of avicularia: small acuminate, small plectrum-shaped, and giant. The distally directed giant avicularia in P. winstonae (Fig. 11F) are very unusual, as giant avicularia in Parasmittina are normally directed proximally, for example in the numerous species described by Soule & Soule (1973), Hayward & Parker (1994) and Harmelin et al. (2009). Harmer (1957: 943) noted the rarity of distally directed giant avicularia when describing material, some from Malaysia and since assigned to P. winstonae by Liu et al. (2001: 802), as Smittina parsevalii (Audouin, 1826). Likewise, the small plectrum-shaped avicularia lateral to the orifice are highly unusual, if not unique, to P. winstonae within Parasmittina. Liu et al. ���s (2001) original description of this species from the South China Sea gives a slightly longer autozooid size (0.37���0.67 mm) than both the Malaysian material described here (0.32���0.45 mm) and specimens from the Solomon Islands (c. 0.45 mm) described by Tilbrook (2006), who did not mention the presence of plectrum-shaped avicularia. This leaves some doubt about whether the three occurrences are truly conspecific, which will require further studies to test., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 19-21, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Liu X., Yin X. & Ma J. 2001. Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing.","Tilbrook K. J. 2006. Cheilostomatous Bryozoa from the Solomon Islands. Santa Barbara Museum of Natural History Monographs 4: 1 - 386.","Soule D. F. & Soule J. D. 1973. Morphology and speciation of Hawaiian and eastern Pacific Smittinidae (Bryozoa, Ectoprocta). Bulletin of the American Museum of Natural History 152: 365 - 440.","Hayward P. J. & Parker S. A. 1994. Notes on some species of Parasmittina Osburn, 1952 (Bryozoa: Cheilostomatida). Zoological Journal of the Linnean Society 110: 53 - 75. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1994. tb 01471. x","Harmelin J. - G., Bitar G. & Zibrowius H. 2009. Smittinidae (Bryozoa, Cheilostomata) from coastal habitats of Lebanon (Mediterranean Sea), including new and non-indigenous species. Zoosystema 31: 163 - 187. http: // dx. doi. org / 10.5252 / z 2009 n 1 a 9","Harmer S. F. 1957. The Polyzoa of the Siboga Expedition, Part 4. Cheilostomata Ascophora II. Siboga Expedition Reports 28 d: 641 - 1147."]}

Cranosina coronata (Hincks, 1881) Fig. 6 Membranipora coronata Hincks, 1881: 147, pl. 10, fig. 1. Cranosina coronata ��� Chimonides & Cook 1994: 44, fig. 1a. Material MALAYSIA: MSL BRY011, Pantai Pasir Hitam, Langkawi, collected intertidally encrusting cobbles from coral reef. Description Colony encrusting, multiserial, unilamellar; growing edge stepped, revealing distolateral pore windows, generally three in each distolateral wall. Ancestrula and early astogeny not observed. Autozooids subhexagonal to almost diamond-shaped, stout, about 0.40���0.50 mm long by 0.34���0.44 mm wide; zooidal boundaries marked by a narrow fissure; opesia occupying most of frontal surface, ovoidal; cryptocyst well developed proximally and laterally, sloping inwards, pustulose, the pustules tending to be radially aligned, especially towards inner side of cryptocyst; gymnocyst sometimes present distolaterally; spines lacking. Ovicells inconspicuous, ooecia small. Intramural buds present. Avicularia present distally of autozooids (Fig. 6C), small, transversely orientated, with long, open-ended rostrum. Remarks According to Tilbrook et al. (2001: p. 45), Cranosina coronata is common throughout the Indo-West Pacific., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 12-13, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Hincks T. 1881. Contributions towards a general history of the marine Polyzoa. IV. Foreign Membraniporina (second series). Annals and Magazine of Natural History, Series 5, 7: 147 - 161. http: // dx. doi. org / 10.1080 / 00222938109459489","Chimonides P. J. & Cook P. L. 1994. Notes on the genus Cranosina (Bryozoa, Cheilostomida). Zoologica Scripta 23: 43 - 49. http: // dx. doi. org / 10.1111 / j. 1463 - 6409.1994. tb 00372. x","Tilbrook K. J., Hayward P. J. & Gordon D. P. 2001. Cheilostomatous Bryozoa from Vanuatu. Zoological Journal of the Linnean Society 131: 35 - 109. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2001. tb 01309. x"]}

Bugula neritina (Linnaeus, 1758) Fig. 8 A���B Sertularia neritina Linnaeus, 1758: 815. Bugula neritina ��� Ryland & Hayward 1977: 162, fig. 78. Material MALAYSIA: MSL BRY014, Batu Uban, Penang, attached to float hanging from fishing jetty. Description Colony erect, weakly mineralized, forming a luxuriant bush comprising bifurcating branches with zooids in two alternating rows opening on one side of branch, purple-brown in colour.Autozooids slender, about 0.48���0.56 mm long by 0.15���0.18 mm wide; opesia occupying almost all of frontal surface, squared-off distally; operculum absent; gymnocyst forming sides and back walls of zooids; spines and cryptocyst lacking; ovicell globular, large and prominent (Fig. 8B), attached to inner distal angle of zooid by a short peduncle. Avicularia lacking. Remarks Considerable attention has been focused on Bugula neritina as a source of the natural product bryostatin. Molecular studies have recently shown that morphologically defined, B. neritina is a complex comprising at least three separate species (Fehlauer-Ale et al. 2014).The complex is very widely distributed in tropical, subtropical and even temperate waters, and is an important fouling taxon dispersed anthropogenically (e.g., Ryland et al. 2011)., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 15-16, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Linnaeus C. 1758. Systema Naturae per Regna tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differetiis, Synonymis, Locis. Edition 10. Laurentii Salvii, Stockholm. Available from http: // biodiversitylibrary. org / page / 726886 [accessed 29 Sep. 2015]","Ryland J. S. & Hayward P. J. 1977. British anascan bryozoans. Cheilostomata, Anasca. Keys and notes for the identification of the species. Synopses of the British Fauna, New Series 10: 1 - 188.","Fehlauer-Ale K. H., Mackie J. A., Lim-Fong G. E., Ale E., Pi M. R. & Waeschenbach A. 2014. Cryptic species in the cosmopolitan Bugula neritina complex (Bryozoa, Cheilostomata). Zoologica Scripta 43: 193 - 205. http: // dx. doi. org / 10.1111 / zsc. 12042","Ryland J. R., Bishop J. D. D., De Blauwe H., El Nagar A, Minchin D., Wood C. A. & Yunnie A. L. E. 2011. Alien species of Bugula (Bryozoa) along the Atlantic coasts of Europe. Aquatic Invasions 6 (1): 17 - 31. http: // dx. doi. org / 10.3391 / ai. 2011.6.1.03"]}

Schizoporella japonica Ortmann, 1890 Fig. 14 A���F Schizoporella unicornsis var. japonica Ortmann, 1890: 49, pl. 3, fig. 35. Schizoporella japonica ��� Ryland et al. 2014: 485, figs 2���5. Material MALAYSIA: MSL BRY019b, BRY024, Kuah jetty, Langkawi, encrusting bivalves fouling a rope hanging from the jetty. Description Colony encrusting, multiserial, unilamellar, growing edge locally developing giant buds (Fig. 14B); vivid yellow-orange when alive (Fig. 14A). Autozooids small, 0.48���0.60 mm long by 0.18���0.38 mm wide, elongate, on average 1.9 �� longer than wide; frontal shield convex, with marginal areolar pores and abundant deep pseudopores, suboral umbo; orifice about as long as wide, 0.11���0.14 mm in both dimensions, sinus wide, shallow, broad condyles occupying outer two-thirds of hingeline on either side of sinus (Fig. 14D); ovicell prominent, porous (Fig. 14 E���F). Adventitious avicularia (Fig. 14D) present in about one-third of autozooids, typically lacking in narrower examples, never more than one per zooid, proximal end level with proximal edge of orifice, directed distolaterally, about 0.11 mm long by 0.06 mm wide; rostrum a high triangle with slightly concave edges and rounded distal end somewhat raised; cross-bar straight; opesia semicircular. Remarks Ryland et al. (2014) comprehensively redescribed Schizoporella japonica and provided information on its geographical distribution, focusing especially on its recent introduction into western Europe. Living colonies of S. japonica from Langkawi have a vibrant yellow-orange colour, similar to the specimen from Friday Harbor, Washington depicted by Ryland et al. (2014: fig. 2d), although some of the colonies described by these authors are redder in colour. The Langkawi material has rather smaller autozooids than is usual for this species, which may reflect the warm ambient seawater temperature, and the avicularia tend to be slightly more laterally orientated., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 25-26, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Ortmann A. 1890. Die Japanische Bryozoenfauna. Bericht uber die von Herrn Dr. L. Doderlein im Jahre 1880 - 81 gemachten Sammlungen. Archiv fur Naturgeschichte 56: 1 - 74. Available from http: // www. biodiversitylibrary. org / item / 30140 page / 9 / mode / 1 up [accessed 29 Sep. 2015]","Ryland J. S., Holt R., Loxton J., Spencer Jones M. E. & Porter J. S. 2014. First occurrence of the nonnative bryozoan Schizoporella japonica Ortmann (1890) in Western Europe. Zootaxa 3780: 481 - 502. http: // dx. doi. org / 10.11646 / zootaxa. 3780.3.3"]}

Sinoflustra amoyensis (Robertson, 1921) Fig. 5 D���F Membranipora amoyensis Robertson, 1921: 49, fig. 6. Sinoflustra amoyensis ��� Liu & Yang 1995: 349, figs 1���2. Material MALAYSIA: MSL BRY010, Batu Muang, Penang, encrusting a barnacle from debris adjacent to a fishing jetty. Description Colony encrusting, multiserial, unilamellar. Ancestrula and early astogeny not observed. Autozooids subrectangular, rounded distally, slender, typically twice as long as wide, 0.43���0.63 mm long by 0.18���0.28 mm wide; zooidal boundaries marked by a narrow fissure; opesia occupying two-thirds of frontal surface, elongate inverted pear-shaped; cryptocyst convex, widest proximally, narrowing distally, covered by small pustules, about 10���16 cryptocystal spinules extending from mural rim over proximal two-thirds of opesia, but not meeting, distal third of opesia broad and spineless (Fig. 5F); gymnocyst lacking. Spines present at distolateral corners of autozooids, spines broken off in studied specimens, leaving a small rounded triangular base with a central pore (Fig. 5F). Avicularia not observed. Remarks The original figure of Robertson (1921: fig. 6) shows zooids with branching spines at the distolateral corners. Although spines are not preserved in the studied material from Penang, their bases are clearly visible in these locations. Robertson���s Chinese material also has slightly shorter cryptocystal spinules and the opesia is ellipsoidal in outline compared with the elongate inverted pear-shaped opesia of the Penang specimens. However, Liu et al. (2001: pl. 18, fig. 3) figured a Chinese specimen of S. amoyensis with longer spinules and opesia that broaden distally. A putative colony of this species epizoic on a turtle has broader autozooids and short cryptocystal spinules (Frazier et al. 1992: fig. 1). In view of the morphological variability encompassed by specimens assigned to the species, the Penang material can be identified as S. amoyensis. Not seen in the Malaysian specimens, but sporadically developed elsewhere in this species, are vicarious avicularia (e.g., Liu et al. 2001: pl. 18, fig. 3). These are longer and wider than the autozooids, and the distal part of the opesia is particularly broad, raised slightly and separated from the proximal part by two small, condyle-like indentations., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 11-12, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Robertson A. 1921. Report on a collection of Bryozoa from the Bay of Bengal and other eastern seas. Records of the Indian Museum 22: 33 - 65. Available from http: // www. biodiversitylibrary. org / item / 41901 page / 71 / mode / 1 up [accessed 29 Sep. 2015]","Liu X. & Yang Z. 1995. Systematic position of Membranipora amoyensis Robertson, 1921 (Membraniporoidea: Cheilostomata). In: Proceedings of the Marine Science Seminar on Taiwan Strait and the Adjacent Seas: 346 - 355. Oceanic Press, Beijing.","Liu X., Yin X. & Ma J. 2001. Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing.","Frazier J. G., Winston J. E. & Ruckdeschel C. A. 1992. Epizoan communities on marine turtles. III. Bryozoa. Bulletin of Marine Science 51: 1 - 8."]}

Adeonella lichenoides (Lamarck, 1816) Fig. 9 A���G Eschara lichenoides Lamarck, 1816: 176. Adeonella lichenoides ��� Hayward 1988: 126, figs 1c, 2���3. Material MALAYSIA: MSL BRY016, Kampung Kuala Temoyong, Langkawi, found among fishing debris. Description Colony erect, rigidly calcified, bushy (Fig. 9A), comprising bifurcating, bifoliate, strap-like branches, about 2.2���2.5 mm wide, with approximately a dozen series of zooids across width of branch.Autozooids small, rounded hexagonal, longer than wide, 0.29���0.38 mm long by 0.19���0.24 mm wide, distinct with deep interzooidal grooves, frontal shield convex, a slight umbo sometimes developed centrally, densely granular in surface texture, with large areolar pores covering most of surface; primary orifice more or less equidimensional, about 0.08 mm long, a shallow U-shaped sinus separated from anter by small condyles (Fig. 9E); secondary orifice somewhat transversely elliptical, a subcircular spiramen separated from secondary orifice by a calcified bridge in mature zooids (Fig. 9 F���G). Gonozooids larger than autozooids, with a broad, shallow orifice separated from a wide spiramen by a deep calcified bridge with a process on proximal edge.Adventitious avicularia (Fig. 9F) developing on frontal shield of autozooids, often laterally to spiramen but occasionally more proximally, generally one or two per autozooid, sometimes absent, orientated variously; rostrum high gothic arch-shaped; cross-bar uncalcified; opesia rounded, a little wider than rostrum. Vicarious avicularia sporadically distributed (Fig. 9D), longer than autozooid, about 0.57 mm long by 0.21 mm wide; rostrum elongate, spatulate with a distal shelf; crossbar uncalcified; opesia roughly semicircular with a sloping cryptocyst-like proximal edge; frontal shield with areolar pores similar to those of autozooids. Kenozooids present along some branch edges. Remarks Found among debris discarded from a fishing net, Adeonella lichenoides is the only rigidly erect bryozoan collected in either Langkawi or Penang during the fieldwork in October 2013. This species is distributed throughout the Indo-West Pacific region (Hayward 1988)., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 17-18, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Lamarck J. B. P. A. de. 1816. Histoire naturelle des Animaux sans Vertebres, Vol. 2. Librairie J. B. Bailliere, Paris.","Hayward P. J. 1988. The Recent species of Adeonella (Bryozoa: Cheilostomata) including descriptions of fifteen new species. Zoological Journal of the Linnean Society 94: 111 - 191. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1988. tb 00105. x"]}

Cradoscrupocellaria sp. Fig. 8 C���F Material MALAYSIA: MSL BRY015, Kuah jetty, Langkawi, fouling a rope hanging from the jetty. Description Colony erect, bushy, jointed, dichotomously branched, ramifications typically after every four zooids, zooids arranged alternately and opening on one side of branch only. Autozooids elongate, about 0.40���0.50 mm long by 0.20���0.24 mm wide; opesia occupying most of frontal surface, oval; gymnocyst moderately well developed proximally; cryptocyst very narrow, smooth; 4 or 5 long, basally articulated spines in distal half of zooid, the one that is more proximal and closest to branch axis forming a scutum bending over frontal membrane and bifid at the end (Fig. 8F); ovicell globular, wider than long, ectooecium with about 4���6 distal pores near distal edge, circular or radially elliptical (Fig. 8F). Avicularia small, located either on: (1) proximal gymnocyst on side closest to branch axis, orientated transversely or proximolaterally and facing outwards, often absent, variable in size, rostrum dentate, tip hooked; or (2) edge of branch close to distal end of autozooids, orientated outwards and facing distally, rostrum triangular, tip hooked. Vibracula located on outer proximal gymnocyst of autozooid, orientated proximally, seta up to 1 mm long. Remarks Vieira et al. (2014) have recently subdivided the species-rich genus Scrupocellaria. The taxonomy of this group is complex and the material collected from Langkawi is tentatively assigned to the genus Cradoscrupocellaria pending a more detailed study., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on page 16, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Vieira L. M., Spencer Jones M. E., Winston J. E., Migotto A. E. & Marques A. C. 2014. Evidence for polyphyly of the genus Scrupocellaria (Bryozoa: Candidae) based on a phylogenetic analysis of morphological characters. PLOS ONE 9 (4): e 95296. http: // dx. doi. org / 10.1371 / journal. pone. 0095296"]}

Acanthodesia cf. falsitenuis (Liu, 1992) comb. nov. Fig. 3 A���C cf. Membranipora falsitenuis Liu, 1992: 112, figs 3���6. Material MALAYSIA: MSL BRY003, Kampung Kuala Temoyong, Langkawi, encrusting an oyster found among fishing debris. Description Colony encrusting, multiserial, unilamellar, locally multilamellar. Ancestrula and early astogenetic stages not observed. Autozooids subhexagonal or subrectangular, distally well-rounded, elongate, 0.53��� 0.73 mm long by 0.30���0.40 mm wide, zooidal boundaries marked by a fissure; mural rim salient, narrow, pustulose, pustules forming a single row; opesia occupying most of frontal surface, rim broadened distally, edged with tiny pustules; gymnocyst lacking; cryptocyst occupying one-third to almost one-half of proximal end of zooid, planar, sparsely pustulose, with a variably developed, anvil-shaped process bearing long spines extending for up to 0.14 mm over opesia (Fig. 3 B���C); no spine bases, spinules or tubercles. Kenozooids developed at convergences between colony lobes (Fig. 3A, upper right corner), variable in shape and size, opesia ovoidal. Remarks The most striking feature of this species is the anvil-shaped, spinose cryptocystal process projecting over the opesia (Fig. 3 B���C). Similar processes were depicted in zooids of two species from the South China Sea ��� Acanthodesia similis (Liu, 1992) (see Liu et al. 2001: pl. 9, fig. 2) and A. falsitenuis (Liu, 1992) (see Liu 1992: fig. 3) ��� and also in Acanthodesia crenulata (Okada, 1923), described by Tilbrook (2006: pl. 2b) from the Solomon Islands. Autozooids of A. similis are about half the length of those of the species described here from Langkawi, but judging from the scale bar shown by Liu (1992) in his figure of A. falsitenuis, this latter species is about the same size as the Langkawi material. However, the cryprocystal process is less developed in the Chinese material of A. falsitenuis and this also has zooids of a more rectangular shape with more ragged edges to the lateral cryptocyst., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 6-8, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Liu X. 1992. On the genus Membranipora (Anasca: Cheilostomata: Bryozoa) from south Chinese seas. Raffles Bulletin of Zoology 40: 103 - 144.","Louis S. & Menon N. R. 2009. Biflustra perambulata n. sp. (Cheilostomata: Bryozoa), a new alien species from Cochin Harbour, Kerala, India. Zootaxa 2066: 59 - 68.","Liu X., Yin X. & Ma J. 2001. Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing.","Tilbrook K. J. 2006. Cheilostomatous Bryozoa from the Solomon Islands. Santa Barbara Museum of Natural History Monographs 4: 1 - 386."]}

Parasmittina raigioidea Liu et al., 2001 Fig. 12 A���C Parasmittina raigioidea Liu et al., 2001: 629, pl. 58, figs 1���2. Material MALAYSIA: MSL BRY020b, Pulau Betong, Penang, encrusting barnacle on oyster raft. Description Colony encrusting, multiserial, unilamellar. Ancestrula and early astogeny not observed. Autozooids subhexagonal, distally rounded, about 0.24���0.51 mm long by 0.19���0.27 mm wide; boundary walls salient with a fine fissure; frontal shield convex, coarsely pustulose, about 20 marginal areolar pores separated by ridges, impinging on central part of shield in older zooids, no pseudopores; primary orifice elliptical, about 0.09 mm long by 0.10 mm wide, lappets developed laterally, lyrula broad, about 9 ��m high by 25 ��m wide at top edge, condyles present about one-quarter along orifice; oral spine bases numbering 1 or 2; ovicell not observed. Avicularia adventitious, variable in size, shape and orientation but not clearly polymorphic (i.e., intermediates existing between common morphologies); usually located lateral to orifice or on proximal frontal shield, the former directed proximally, the latter distally or laterally, 0.08���0.23 mm long by 0.05���0.08 mm wide; rostrum acutely triangular to spatulate with a moderate shelf, straight or slightly curved; cross-bar calcified, straight to concave; opesia semielliptical. Remarks Acknowledging the problems in discriminating between species of Parasmittina, this material is tentatively assigned to P. raigioidea, a species described originally from South China, on account of the similar range of avicularian morphologies to those seen in pl. 58, figs 1 and 2 of Liu et al. (2001). These authors recognised two types of small avicularia ��� triangular and spatulate ��� as well as giant avicularia. The Malaysian material shows a range of avicularian morphologies and sizes and it is unclear whether there is one highly variable type of avicularium or several polymorphs., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 21-23, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Liu X., Yin X. & Ma J. 2001. Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing."]}

Scorpiodinipora costulata (Canu & Bassler, 1929) Fig. 15 G���I Schizoporella costulata Canu & Bassler, 1929: 317, pl. 36, fig. 10. Scorpiodinipora costulata ��� Harmelin et al. 2012: 129, figs 1���4. Material MALAYSIA: MSL BRY012c, Kampung Kuala Temoyong, Langkawi, colony encrusting interior of large gastropod shell. MSL BRY025, Pulau Betong, Penang, on gastropod from oyster rafts. Description Colony encrusting, multiserial, unilamellar; buttressed pore chambers visible at stepped growing edge. Ancestrula and early astogeny not observed. Autozooids rounded subhexagonal, 0.35���0.41 mm long by 0.23���0.26 mm wide, zooidal boundaries with narrow grooves; frontal shield convex, mammillated, bordered by large areolar pores separated by radial ridges, imperforate centrally; orifice large relative to zooid size, ovoidal, longer than wide, 0.13���0.15 mm long by 0.09���0.10 mm wide, anter and poster similarly sized, well rounded, separated by condyles that mark beginning of a rim outlining anter; oral spines lacking; ovicells absent. No avicularia. Remarks This widely distributed species was revised by Harmelin et al. (2012) who noted its preference for gastropod shells. The two colonies collected in Malaysia, one from Langkawi and the other from Penang, are both on gastropod shells, though not a living gastropod in the case of the Langkawi specimen, which encrusts an interior surface., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on page 29, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Canu F. & Bassler R. S. 1929. Bryozoa of the Philippine region. United States National Museum Bulletin 100 (9): 1 - 685. Available from http: // www. biodiversitylibrary. org / item / 32602 page / 5 / mode / 1 up [accessed 29 Sep. 2015]","Harmelin J. - G., Vieira L. M., Ostrovsky A. N., Caceres-Chamizo J. P. & Sanner J. 2012. Scorpiodinipora costulata (Canu & Bassler, 1929) (Bryozoa, Cheilostomata), a taxonomic and biogeographic dilemma: complex of cryptic species or humanmediated cosmopolitan colonizer? Zoosystema 34: 123 - 138. http: // dx. doi. org / 10.5252 / z 2012 n 1 a 5"]}

Parasmittina sp. 2 Fig. 12 G���I Material MALAYSIA: MSL BRY012b, Kampung Kuala Temoyong, Langkawi, colony encrusting interior of large gastropod shell. Description Colony encrusting, multiserial, unilamellar. Ancestrula and early astogeny not observed. Autozooids subhexagonal, small, about 0.24���0.32 mm long by 0.18���0.24 mm wide; boundary walls salient with a fine fissure; frontal shield convex, coarsely pustulose, marginal areolar pores large, no pseudopores; orifice slightly wider than high, about 0.09 mm long by 0.10 mm wide, lappets well-developed, highest proximolaterally, tapering distally, a sloping shelf-like proximal orificial rim extending between lappets, lyrula variable, condyles stout; usually two oral spine bases, occasionally one or three; ovicell broader than long, observed only partly formed.Avicularia adventitious present on a small minority of autozooids, located proximolaterally or laterally of orifice, directed proximally or less often laterally, variable in size, up to 0.21 mm long by 0.08 mm wide; rostrum acutely triangular with rounded end, shelf variable, occupying up to one-half length; cross-bar complete or with a medial gap; opesia semi-elliptical. Remarks Compared with the other three species of Parasmittina, this unidentified species has very sparse avicularia and prominent lappets highest at the proximolateral corners of the orifice and tapering in height distally (Fig. 12I). There are some similarities with Parasmittina hastingsae Soule & Soule, 1973 (see Tilbrook et al. 2001: fig. 14c���d), but the latter has larger autozooids with long condyles and the lappets are highest mediolaterally rather than proximolaterally., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 23-24, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Soule D. F. & Soule J. D. 1973. Morphology and speciation of Hawaiian and eastern Pacific Smittinidae (Bryozoa, Ectoprocta). Bulletin of the American Museum of Natural History 152: 365 - 440.","Tilbrook K. J., Hayward P. J. & Gordon D. P. 2001. Cheilostomatous Bryozoa from Vanuatu. Zoological Journal of the Linnean Society 131: 35 - 109. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2001. tb 01309. x"]}

Hippoporina indica Pillai, 1978 Fig. 13 A���F Hippoporina indica Pillai, 1978: 62, figs 1���4. Hippoporina indica ��� McCann et al. 2007: 331, fig. 7a���d. Material MALAYSIA: MSL BRY017b, Kuah jetty, Langkawi, fouling mussel shell attached to rope hanging from jetty. MSL BRY022, Pulau Betong, Penang, fouling a bivalve from an oyster raft. MSL BRY023, Sungai Menghulu, Langkawi, fouling a barrel. Description Colony encrusting, multiserial, unilamellar, except for frontal buds covering early astogenetic stages and apparent reparative growths. Autozooids subrectangular, elongate, 0.35���0.63 mm long by 0.23���0.33 mm wide; frontal shield gently convex, pustulose, porous, with large marginal areolar pores and large pseudopores, which are lacking from an apron proximal of orifice; orifice longer than wide (Fig. 13B), about 0.13 mm long by 0.11 mm wide, sinus broad and shallow, with medial edge almost straight, a pair of proximally directed, pointed condyles separating sinus from semicircular poster, closed by cryptocystal calcification in some zooids (Fig. 13C); ovicell hyperstomial, prominent, broader than long, about 0.16���0.18 mm long by 0.21���0.23 mm wide; about 10���20 rimmed pores of various shapes and sizes, becoming overgrown from the margins by a lamina of interior wall (Fig. 13 D���E). Avicularia adventitious, small, about 0.10 mm long by 0.07 mm wide, normally located laterally to orifice and directed distolaterally towards orifice, usually single, lacking in many zooids, occasional avicularia with variable orientations present more proximally; rostrum pointed, arch-shaped; cross-bar calcified, narrow; opesia semielliptical, broader than long. Remarks Despite being described as late as 1978 (from Bombay Harbour), Hippoporina indica is rapidly becoming widespread as an invasive fouling species. It has been reported from the southeastern USA (McCann et al. 2007), New Zealand (Gordon et al. 2008) and Australia (Tilbrook 2012), and its presence in Penang and Langkawi is therefore unsurprising., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 24-25, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Pillai S. R. M. 1978. A new species of Hippoporina (Ectoprocta, Ascophora) from Bombay coast. Current Science (Bangalore) 47: 61 - 63.","McCann L. D., Hitchcock N. G., Winston J. E. & Ruiz G. M. 2007. Non-native bryozoans in coastal embayments of the southern United States: new records for the western Atlantic. Bulletin of Marine Science 80: 319 - 342.","Gordon D. P., Hosie A. M. & Carter M. C. 2008. Post- 2000 detection of warm-water alien bryozoan species in New Zealand - the significance of recreational vessels. Virginia Museum of Natural History, Special Publication 15: 37 - 48.","Tilbrook K. J. 2012. Bryozoa, Cheilostomata: First records of two invasive species in Australia and the northerly range extension for a third. Check List 8 (1): 181 - 183."]}

Acanthodesia perambulata (Louis & Menon, 2009) comb. nov. Fig. 3 D���F Biflustra perambulata Louis & Menon, 2009: 61, figs 2���7. Material MALAYSIA: MSL BRY004, Pantai Pasir Hitam, Langkawi, on floating plastic washed ashore. Description Colony encrusting, multiserial, unilamellar, locally multilamellar (Fig. 3D). Ancestrula and early astogenetic stages not observed. Autozooids rounded rectangular, gently convex distally, 0.43���0.60 mm long by 0.30���0.50 mm wide, the first zooids in new rows narrow and about 2 �� longer than wide, later zooids becoming progressively broader until approximately equidimensional immediately before row bifurcation (Fig. 3F); zooidal boundaries marked by a fine fissure; mural rim salient, narrow, pustulose; opesia occupying nearly all frontal surface, ovoidal; gymnocyst lacking; cryptocyst narrow, broadest at proximolateral corners, planar, with sporadic pustules that increase in abundance and become dense around edge of opesia; no spine bases, spinules or tubercles. Kenozooids developed at convergences between colony lobes, variable in shape and size, opesia ovoidal. Remarks This species was originally described from Cochin, India, where colonies are erect and bilamellar, unlike the encrusting colonies from Langkawi assigned here to Acanthodesia perambulata. However, the skeletal morphology and dimensions of the autozooids is almost identical in the Indian and Malaysian material and it is known that a single species can exist as either encrusting or erect bifoliate colonies. Tilbrook & Gordon (2015) list this species (as Biflustra perambulata) from the Straits of Johor, Singapore. Compared to the similar species Acanthodesia grandicella Canu & Bassler, 1929 (e.g., Gordon et al. 2008; Tilbrook 2012), A. perambulata has smaller zooids that are less granular and have weaker calcification. Nevertheless, molecular sequence data is needed to show that these fouling invasive species really are distinct from one another., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on page 8, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Louis S. & Menon N. R. 2009. Biflustra perambulata n. sp. (Cheilostomata: Bryozoa), a new alien species from Cochin Harbour, Kerala, India. Zootaxa 2066: 59 - 68.","Tilbrook K. J. & Gordon D. P. 2015. Bryozoa from the Straits of Johor, Singapore, with the description of a new species. Raffles Bulletin of Zoology, Supplement 31: 255 - 263.","Canu F. & Bassler R. S. 1929. Bryozoa of the Philippine region. United States National Museum Bulletin 100 (9): 1 - 685. Available from http: // www. biodiversitylibrary. org / item / 32602 page / 5 / mode / 1 up [accessed 29 Sep. 2015]","Gordon D. P., Hosie A. M. & Carter M. C. 2008. Post- 2000 detection of warm-water alien bryozoan species in New Zealand - the significance of recreational vessels. Virginia Museum of Natural History, Special Publication 15: 37 - 48.","Tilbrook K. J. 2012. Bryozoa, Cheilostomata: First records of two invasive species in Australia and the northerly range extension for a third. Check List 8 (1): 181 - 183."]}

Microporella sp. Fig. 15 D���G Material MALAYSIA: MSL BRY012d, Kampung Kuala Temoyong, Langkawi, colony encrusting interior of large gastropod shell. Description Colony encrusting, multiserial, unilamellar. Ancestrula and early astogeny not observed. Autozooids small, about 0.35���0.37 mm long by 0.28 mm wide, subhexagonal, rounded distally; distinct with interzooidal grooves; frontal shield convex, porous, a few elongate areolar pores around perimeter and numerous smaller circular pseudopores elsewhere, coarsely pustulose; orifice semi-elliptical, small, about 0.06 mm long by 0.08���0.09 mm wide, proximal edge straight, without teeth or condyles, distal edge beaded; oral spines numbering three in non-ovicellate zooids, one distal medial and two lateral, level with orifice mid-length, not visible in ovicellate zooids; ascopore crescent-shaped, about 0.04 mm wide, toothed, set in a rugose prominence about 0.06 mm long by 0.08 mm wide, close to proximal edge of orifice, but separated by a narrow band of non-porous frontal shield (Fig. 15E); ovicell personate, proximal border of secondary orifice broadly U-shaped, ooecium broader than long, about 0.16 mm long by 0.20 mm wide, porous except in the most proximal medial part, calcification continuous with that of frontal shield of distal zooid, on which ovicell rests without extending as far distally as ascopore. Adventitious avicularia present in a minority of autozooids, single, located about mid-length on autozooid, directed laterally outwards, small, about 0.08 mm long by 0.06 mm wide; rostrum with concave sides and open end (i.e., channeled); mandible setose, almost twice length of rostrum; opesia semicircular. Remarks Microporella is a highly species-rich genus with a global distribution (Taylor & Mawatari 2005). Differences between species can be very subtle. The species described here is characterized by having: (1) unpaired avicularia, often lacking altogether, laterally directed; (2) a broad crescent-shaped ascopore set in a wide rugose prominence; (3) three oral spines; and (4) an orifice lacking both teeth and condyles along the proximal edge but having a beaded distal edge. A beaded (or denticulate) distal orificial edge occurs in relatively few species of Microporella, such as M. serrata Mawatari & Suwa, 1998, M. rogickae Winston, Hayward & Craig, 2000, and M. browni Harmelin, Ostrovsky, C��ceres-Chamizo & Sanner, 2011. In M. serrata the avicularia are located much more distally than in the species from Langkawi and the ascopore is cribriform. Condyles are present at the proximolateral corners of the orifice in both this species and M. rogickae, the latter having 4���5 oral spines (cf. 3 in the species described here) and a much thicker proximal edge to the personate ovicells, whereas in M. serrata 4 oral spines are present and the ovicells are not personate. Although the Langkawi species may be new, its formal description must await additional material., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on page 28, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Taylor P. D. & Mawatari S. F. 2005. Preliminary overview of the cheilostome bryozoan Microporella. In: Moyano G. H. I., Cancino J. M. & Wyse Jackson P. N. (eds) Bryozoan Studies 2004: 329 - 339. Balkema, Leiden."]}

Acanthodesia Canu & Bassler, 1919 Remarks The genus Acanthodesia, with the type species Membranipora savartii Audouin, 1826 (see Taylor & Foster 1994), is here interpreted to include numerous species once assigned to Membranipora but having more robustly calcified colonies, or, alternatively, to Biflustra but generally having encrusting colonies rather than the erect vincularian colonies of the type species of this genus. Although many bryozoologists use Biflustra in preference to Acanthodesia, the critical early astogeny is unknown in the type species of the former genus. The paucity of skeletal morphological characters ��� avicularia and ovicells are lacking ��� makes species identification difficult. Molecular studies are much needed in this genus to clarify its diversity., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on page 6, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Taylor P. D. & Foster T. S. 1994. Bryozoa from the Plio-Pleistocene of Tobago, West Indies. Tertiary Research 15: 1 - 16."]}

Antropora minor (Hincks, 1880) Fig. 7A Membranipora trifolium var. minor Hincks, 1880: 87, pl. 11, fig. 6. Antropora minor ��� Tilbrook 1998: 34, fig. 2a���f. Material MALAYSIA: MSL BRY012a, Kampung Kuala Temoyong, Langkawi, colony encrusting interior of large gastropod shell. Description Colony encrusting, multiserial, unilamellar or multilamellar. Ancestrula and early astogeny not observed. Autozooids subovoidal, about twice as long as wide, 0.34���0.40 mm long by 0.18���0.40 mm wide; zooidal boundaries marked by a narrow fissure; opesia occupying one-half to two-thirds of frontal surface, pear-shaped, generally pinched slightly at about two-thirds of length; cryptocyst well developed proximally and shelf-like, tapering distally, pustulose; gymnocyst visible proximally and proximolaterally, narrow, tapering distally; spines lacking. Ovicells not observed in studied Malaysian material but elsewhere described as small, partly immersed in distal zooid and with cap-like ooecia (e.g., Tilbrook 1998). Avicularia interzooidal, directed distally, small, about 0.06���0.09 mm long by 0.04���0.05 mm wide, present at proximal corner or less often at two proximolateral corners of most autozooids; rostrum rounded, a little longer but narrower than opesia. Remarks Large vicarious avicularia sometimes develop in this species (e.g., Tilbrook 1998: fig. 2e) but were not seen in the studied colony from Langkawi. Antropora minor is widely distributed in the tropics, occurring in the Caribbean, South Atlantic, Pacific and Indian Oceans, including East Sumbawa, Malaysia (Tilbrook 1998)., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 13-14, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Hincks T. 1880. Contributions towards a general history of the marine Polyzoa. Annals and Magazine of Natural History, Series 5, 6: 69 - 92. http: // dx. doi. org / 10.1080 / 00222938009458895","Tilbrook K. J. 1998. The species of Antropora Norman, 1903 (Bryozoa: Cheilostomatida), with the description of a new genus in the Calloporoidea. Records of the South Australian Museum 31: 25 - 49. Available from http: // www. biodiversitylibrary. org / item / 126052 page / 28 / mode / 1 up [accessed 29 Sep. 2015]","Hincks T. 1881. Contributions towards a general history of the marine Polyzoa. IV. Foreign Membraniporina (second series). Annals and Magazine of Natural History, Series 5, 7: 147 - 161. http: // dx. doi. org / 10.1080 / 00222938109459489"]}

Nellia oculata Busk, 1852 Fig. 7 B���E Nellia oculata Busk, 1852: 18, pl 64, fig. 6, pl 65 bis, fig. 5. Nellia oculata ��� Winston et al. 2014: 161, fig. 13. Material MALAYSIA: MSL BRY013, Balik Pulau, Penang, small colony washed ashore on a sandy beach. Description Colony erect, articulated, forming a tiny dichotomously branched bush composed of quadriserial internodes with 4���7 autozooids each linked by chitinous joints, basally rooted by kenozooids.Autozooids slender, about 0.38���0.50 mm long by 0.13���0.15 mm wide; opesia occupying about three-quarters of surface, elongate elliptical in shape; gymnocyst widest proximally, tapering distally along sides of zooid, spines lacking; cryptocyst forming a slightly sunken, inwardly sloping crescent at proximal end of opesia, smooth-surfaced; various pores and depressions visible through opesia in basal walls of zooids, plus a low spinose, curved ridge on distal interior wall of zooid (Fig. 7C); ovicell not observed in studied material but described elsewhere as endozooidal (Winston et al. 2014). Avicularia tiny (Fig. 7E), located as pairs on left and right proximolateral gymnocyst of autozooids, inclined to plane of cryptocyst and directed proximally; rostrum rounded; cross-bar calcified, straight; opesia smaller than rostrum, with a slightly depressed cryptocyst; countersunk pore positioned on autozooidal gymnocyst beneath avicularium. Remarks This species has often been regarded as a junior synonym of Nellia tenella (Lamarck, 1816), but following Winston et al. (2014) we here retain Busk���s species name oculata pending a revision of Lamouroux���s material to test the synonymy. Ostensibly, Nellia oculata has a wide tropical and subtropical distribution, as well as a long fossil record, but there is need for a taxonomic re-evaluation using molecular data., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 14-15, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Busk G. 1852. Catalogue of Marine Polyzoa in the Collection of the British Museum. Part I. Cheilostomata (part). Trustees of the British Museum, London.","Winston J. E., Vieira L. M. & Woollacott R. M. 2014. Scientific results of the Hassler Expedition. Bryozoa. No. 2. Bulletin of the Museum of Comparative Zoology 161 (5): 139 - 239.","Hincks T. 1880. Contributions towards a general history of the marine Polyzoa. Annals and Magazine of Natural History, Series 5, 6: 69 - 92. http: // dx. doi. org / 10.1080 / 00222938009458895","Lamarck J. B. P. A. de. 1816. Histoire naturelle des Animaux sans Vertebres, Vol. 2. Librairie J. B. Bailliere, Paris."]}

Aetea ligulata Busk, 1852 Fig. 2 A���F Aetea ligulata Busk, 1852: 31, pl. 42. Aetea ligulata ��� Osburn 1950: 13, pl. 1, fig. 4. Material MALAYSIA: MSL BRY001a, Pantai Pasir Hitam, Langkawi, collected intertidally from coral reef, fouling Hippopodina feegeensis. MSL BRY002, Kampung Kuala Temoyong, Langkawi, encrusting shell found among fishing debris. Description Colony encrusting, uniserial, runner-like with widely spaced branches (Fig. 2 A���B), delicate, feebly calcified. Autozooids slender with elongate pyriform proximal base and an erect distal part; proximal base 1.1���1.3 mm long by about 0.13 mm maximum width (Fig. 2C), with irregular transverse growth wrinkles and locally minutely porous or with finely reticulate ornamentation, a narrow proximal cauda of about same length as broader distal part; erect part often broken off or collapsed, about 0.42 mm long by 0.07 mm maximum width, proximally with widely spaced, hoop-like annulations about 0.02 mm apart (Fig. 2D), expanding a little distally where an elongated opesia, 0.25 mm long by 0.05 mm wide, is developed on proximal-facing side (Fig. 2E). Remarks This species was erected by Busk (1852) based on material collected by Charles Darwin from the coast of Patagonia and the Magellan Strait. Busk���s figure shows clearly the coarsely annulated erect parts of the zooids, very different from the finely striated annulations seen in the two common cosmopolitan species Aetea anguina (Linnaeus, 1758) and A. sica (Couch, 1844). A modern systematic study of Aetea combining morphological and molecular data is required and in the meantime there must be reservations about the identities of species with such seemingly wide latitudinal distributions as A. ligulata. Abundant evidence of damage and repair can be observed in the skeleton. The skeletons of some zooids were apparently partly destroyed, eliciting reparative growth, sometimes on multiple occasions (Fig. 2C). Broken zooids may also reveal internal stolon-like structures (Fig. 2F), indicating growth through the dead zooid to re-establish connections between zooids and allow growth from the open ends of branches. Although the straggly, runner-like form of Aetea and various other uniserial cheilostomes would suggest a reduced commitment to maintaining the integrity of the colony, reparative growth that renews links between zooids and re-uses substrate space once occupied by dead zooids is a common feature of such bryozoans, Recent and fossil (Taylor 1988)., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 5-6, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Busk G. 1852. Catalogue of Marine Polyzoa in the Collection of the British Museum. Part I. Cheilostomata (part). Trustees of the British Museum, London.","Osburn R. C. 1950. Bryozoa of the Pacific coast of America. Part 1, Cheilostomata-Anasca. Allan Hancock Pacific Expeditions 14: 1 - 269. Available from http: // www. biodiversitylibrary. org / item / 41904 page / 13 / mode / 1 up [accessed 29 Sep. 2015]","Linnaeus C. 1758. Systema Naturae per Regna tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differetiis, Synonymis, Locis. Edition 10. Laurentii Salvii, Stockholm. Available from http: // biodiversitylibrary. org / page / 726886 [accessed 29 Sep. 2015]","Taylor P. D. 1988. Colony growth pattern and astogenetic gradients in the Cretaceous cheilostome bryozoan Herpetopora. Palaeontology 31: 519 - 549."]}

Jellyella eburnea (Hincks, 1891) Fig. 4 Membranipora eburnea Hincks, 1891: 289, pl. 7, fig. 5. Jellyella eburnea ��� Taylor & Monks 1997: 42, figs 1���2, 4���13, 16���19. Material MALAYSIA: MSL BRY008, Pantai Chenang, Langkawi, on plastic and glass bottles. Description Colony encrusting, multiserial, unilamellar (Fig. 4A, E). Ancestrula twinned. Autozooids subhexagonal, rounded distally, twice as long as wide, 0.52���0.67 mm long by 0.29���0.33 mm wide; zooidal boundaries marked by a fine fissure; opesia occupying nearly all frontal surface, roughly ovoidal; cryptocyst forming a narrow, crescent-shaped shelf at distal end of zooid (Fig. 4B); gymnocyst variably developed, when well-developed folded into rucks or spines projecting over proximal and later margins of opesia (Fig. 4C); basal walls with large uncalcified window; intramural buds present but rare. Remarks This species was fully described by Taylor & Monks (1997) who made it the type species of the new genus Jellyella, noting the pan-oceanic, tropical to subtropical occurrence of J. eburnea as an encruster of floating shells of the squid Spirula, and occasionally of the living planktonic gastropod Janthina. The pseudoplanktonic ecology of J. eburnea has apparently ���pre-adapted��� it to the colonization of floating anthropogenic debris, especially plastic objects (e.g., Moyano 2005) but also glass bottles (Fig. 4 D���E)., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 9-10, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Hincks T. 1891. Contributions towards a general history of the marine Polyzoa. XV. South-African and other Polyzoa. Annals and Magazine of Natural History, Series 6, 7: 285 - 298. http: // dx. doi. org / 10.1080 / 00222939109460610","Taylor P. D. & Monks N. 1997. A new cheilostome bryozoan genus pseudoplanktonic on molluscs and algae. Invertebrate Biology 116: 39 - 51.","Moyano H. 2005. Bryozoa de la Expedicion Chilena CIMAR 5 Islas Oceanicas I: el genero Jellyella Taylor & Monks 1997 (Bryozoa, Cheilostomatida) en la Isla de Pascua. Ciencia y Tecnologia de Mar 28 (2): 87 - 90."]}

Hippopodina feegeensis (Busk, 1884) Fig. 15 A���C Lepralia feegeensis Busk, 1884: 144, pl. 22, fig. 9, 9a, 9b. Hippopodina feegeensis ��� Tilbrook 1999: 451, fig. 1a���h. Material MALAYSIA: MSL BRY001b, Pantai Pasir Hitam, Langkawi, collected intertidally from coral reef. Description Colony encrusting, multiserial, unilamellar or multilamellar, often large in size; growing edge revealing line of buttressed pores in transverse walls. Ancestrula and early astogeny not observed in studied Malaysian material; elsewhere ancestrula comprising a tetrad of zooids (e.g., Tilbrook 1999: fig. 1b). Autozooids large, 0.75���1.00 mm long by 0.42���0.58 mm wide, subrectangular to subhexagonal; distinct with interzooidal grooves; frontal shield gently convex, perforated by numerous small, closely spaced pores, pustulose; orifice large, hoof-shaped, almost equidimensional, 0.20���0.22 mm long by 0.18���0.20 mm wide, proximal edge gently concave, wide, condyles rounded, lateral; ovicell large (Fig. 15B), 0.46���0.50 mm long by 0.44���0.50 mm wide, evenly porous, calcification resembling frontal shield but lacking pustules, primary orifice of ovicellate zooids a little larger than in non-ovicellate zooids, secondary orifice broad, 0.12 mm long by 0.24 mm wide, incipient ovicell evident as a sparsely porous, smooth depression in proximal gymnocyst of distal zooid. Adventitious avicularia (Fig. 15C) present in less than one-half of autozooids, singly or paired, 0.18���0.26 mm long by 0.12���0.16 mm wide; located distolaterally of orifice, directed transversely towards midline of supporting autozooid; rostrum a high triangle with slightly concave sides, pointed distally; cross-bar calcified in most, straight; opesia semicircular. Remarks This is a common species circumtropically, often occurring in coral reefs, which is widely distributed in the Pacific, Indian and Atlantic Oceans and also recorded from the Mediterranean and Red Seas (see Powell 1969: fig. 2). The large sizes of both the colonies and their constituent zooids are notable. Colonies of this species found at Pantai Pasir Hitam, Langkawi were remarkable for being concentrated in a small area and absent from similar-looking habitats in the area. This suggests localized recruitment, perhaps from a single founding colony., Published as part of Taylor, Paul D. & Tan, Shau-Hwai Aileen, 2015, Cheilostome Bryozoa from Penang and Langkawi, Malaysia, pp. 1-34 in European Journal of Taxonomy 149 on pages 26-28, DOI: 10.5852/ejt.2015.149, http://zenodo.org/record/3787554, {"references":["Busk G. 1884. Report on the Polyzoa collected by H. M. S. Challenger during the years 1873 - 1876. Part 1. The Cheilostomata. Report on the Scientific Results of the Voyage of the H. M. S. \" Challenger \", Zoology 10 (30): 1 - 216.","Tilbrook K. J. 1999. Description of Hippopodina feegeensis and three other species of Hippopodina Levinsen, 1909 (Bryozoa: Cheilostomatida). Journal of Zoology 247: 449 - 456. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1999. tb 01008. x","Ortmann A. 1890. Die Japanische Bryozoenfauna. Bericht uber die von Herrn Dr. L. Doderlein im Jahre 1880 - 81 gemachten Sammlungen. Archiv fur Naturgeschichte 56: 1 - 74. Available from http: // www. biodiversitylibrary. org / item / 30140 page / 9 / mode / 1 up [accessed 29 Sep. 2015]","Canu F. & Bassler R. S. 1929. Bryozoa of the Philippine region. United States National Museum Bulletin 100 (9): 1 - 685. Available from http: // www. biodiversitylibrary. org / item / 32602 page / 5 / mode / 1 up [accessed 29 Sep. 2015]","Powell N. A. 1969. Indo-Pacific Bryozoa new to the Mediterranean coast of Israel. Israel Journal of Zoology 18: 157 - 168. http: // dx. doi. org / 10.1080 / 00212210.1969.10688281"]}

Glyphidrilus kotatinggi Chanabun & Panha, sp. n. (Figs 1, 2B, 4, Table 1) Type material. Holotype: One clitellate (CUMZ 3274) (Fig. 4), Kota Tinggi waterfall, Johor, Malaysia, 62 meters in elevation, S. Panha, R. Chanabun, P. Tongkerd and C. Sutcharit leg., 20 May 2011. Four paratypes: 3 clitellates (CUMZ 3275), 1 clitellate (ZMH 14574), same collection data as for holotype. Type locality. River banks of Kota Tinggi waterfall, Johor, Malaysia, 01°49'44.5''N, 103°50'4.1"E, 62 m asl. Etymology. This new species was named after the Kota Tinggi waterfall, a famous waterfall. This is the first record of Glyphidrilus from this place. Diagnosis. Medium sized limicolous earthworm with distinct expanded tissues of clitellar wing organs on the lateral sides of the body in 20, 21–¼26, 26. Clitellum in 17, 18–28, 29, 30. Female pores, male pores not visible. Inseminated spermathecae visible through body wall in 14/15–16/17. Genital markings post-setal median unpaired without rim on aa in 17, 19, post-setal lateral paired or asymmetrically distributed without rim near b or bc in 11–15, 17, 18, 19, 20 and 26–27. Four pairs of seminal vesicles in 9–12, with the pair in segment 12 largest. Three paired of hearts in 9–11. Intestinal origin in 14. Ovaries in 13–14. Prostate and accessory glands absent. Spermathecae in 13/14–16/17, one to four on each side per segment. Description of holotype. Dimensions: body length 195 + mm, diameter 3.1 mm in segment 8, 3.1 mm before clitellar wings in segment 20, 3.2 mm behind wings in segment 27 in intraclitellar region. Body cylindrical in anterior part, quadrangular in transverse section behind clitellum. 284 + segments. Body colour pale brown with variations from red to pink at adjacent tissues of wing portion in different individuals of newly collected specimens after placement in 30% ethanol for narcotization. Dorsal surface considerably broader than the ventral at posterior end. Clitellar wings on ventro-lateral part of clitellum in 21–¼26, 5.4 mm long, about 0.6 mm wide on both sides. Prostomium zygolobous. Dorsal pores absent. Clitellum annular in 18–29. Four pairs of setae per segment from 2, setal formula aa:ab:bc:cd:dd = 0.9:0.4:1.6:0.5:1.5 in segment 8. Female pores and male pores not visible. Inseminated spermathecae visible through body wall on b, c, and d line in 14/15–16/17. Genital markings: postsetal median unpaired without rim on aa in 19, post-setal lateral paired without rim near b in 11–15, 20 and 26–27; in 11–13 three genital markings on each side, in 14–15 only one. Septa 4/5–7/8 thicker, 8/9–11/12 thick and 12/13 to the last segment thin. Gizzard small, globular within 8–9. Intestine enlarged from 14. Dorsal blood vessel aborted anterior to 9. Hearts from 9–11. No nephridia distinguishable in first eleven segments. Four pairs of seminal vesicles in 9–12, with pair in segment 12 larger than the others. Ovaries in 13–14. Prostate and accessory glands absent. Spermathecae in 13/14–16/17, three to four on each side in 14/15–16/17, inseminated, one or two on each side in 13/14, not inseminated. Number of spermathecae on the right side: two in 13/14, three in 14/15, four in 15/16 and in 16/17, on the left side: one in 13/14, three in 14/ 15 and 16/17, and four in 15/16. Variation. Body length of the four paratypes 151 + – 187 mm, with 221 + –415 segments. Wings in 20, 21–¼26, 26 (covering 6–7 segments), clitellum in 17, 18–28, 29, 30 (covering 11–14 segments). Genital markings: postsetal median unpaired without rim on aa in 17, 19, post-setal lateral paired or asymmetry without rim near b or bc in 11–15, 17, 18, 19, 20 and 26–27. Habitat. Found on the shore but in proximity to the river water, in the loamy sand topsoil with 82% sand, 14.28% silt and 3.57% clay, with pH 7.54, and also under the water at about 15–25 cm in depth. The river bank near the waterfall was covered with worm casts. Distribution. The new species is known only from the type locality. Remarks. Five described species from areas near Malaysia include G. kukenthali, G. malayanus, G. gatesi, G. singaporensis, and G. vangviengensis. They are different from the new species with respect to spermathecae, genital markings, clitellum and wing locations. Glyphidrilus kotatinggi sp. n. is most similar to G. singaporensis Shen & Yeo, 2005, described from a locality nearby (only 45 km apart). Referring to the original description of the latter, G. kotatinggi differs in larger body size (115 + –195 + mm vs. 112–142 mm), higher segment number (221 + –415 + vs. 163–220), and in the absence of unpaired post-wing genital markings. Further possible differences (clitellum covering 11–14 segments and wings covering 6–7 segments vs. clitellum covering 9–14 segments and wings covering 5–6 segments) are inconclusive due to intraspecific variations in both species that lead to character overlap. For a detailed comparison, we re-investigated the holotype of G. singaporensis and we collected new topotypic material. As a result, we confirm the three above-mentioned differences of G. kotatinggi to G. singaporensis. A comparison of G. kotatinggi with G. singaporensis (specified for holotype, paratypes, and topotypic material) is given in Table 1. Differences of further species to G. kotatinggi sp. n. are as follows (see also Table 1): G. kukenthali from Sarawak differs in slightly longer wings in 18, ½18, 19–24, ½24, longer clitellum in ½13–34, hearts in 7–11, and spermathecae in 14–18. G. malayanus from the Malay Peninsula has a bit shorter wings in 17, ¾18, 18–⅓21, ⅓22, 22, a shorter clitellum in 15–25, 26, intestinal origin in 16, and spermathecae in 14/15–16/17. G. gatesi from Johor, Malaysia has wings in 18, ½19, 19–½24, 24, a shorter clitellum, in ½17, 17, 18–25, ½26, 26, intestinal origin in 18, and spermathecae in 15–17. G. vangviengensis reported from the river bank of Song River at Vangvieng, Laos, has longer wings in 24, 25–31, 32, longer clitellum in 19, 20–35, 36, 37, intestinal origin in 16, and spermathecae are absent. G. bisegmentus sp. n. from Air Banun Pandig, Perak, has shorter wings, shorter clitellum, and genital markings and spermathecae are absent. G. peninsularis sp. n. from Sungei Bantang, has longer wings, a slighlty longer clitellum, four pairs of hearts in 8–11, and spermathecae in 14/15–17/18. Material of other species examined for comparison. Glyphidrilus kukenthali Michaelsen, 1896: 195, pl. 13, Fig. 1; type locality: Borneo, Barem River, Sarawak. Glyphidrilus malayanus Michaelsen, 1902: 35; one clitellate (syntype ZMH V 5875); type locality: Malay Peninsula, Lubock Paku, Pahang River. Glyphidrilus gatesi Shen & Yeo, 2005: 16, Fig. 1; one clitellate (holotype ZRC 1974.12.2.51) and 9 clitellates and 2 aclitellates (paratypes ZRC 1974.12.2.52–62); type locality: Sungei Kayu, swamp forest near River Sedili, Johor, Malaysia. Glyphidrilus singaporensis Shen & Yeo, 2005: 18, Fig. 3; one clitellate (holotype ZRC) and 2 clitellates and 9 aclitellates (paratypes ZRC) and 10 clitellates and 9 aclitellates (topotypes CUMZ 3273); type locality: Jungle Fall Valley, Bukit Timah, Singapore. Glyphidrilus vangviengensis Panha & Chanabun, 2011: 216, Figs. 1–3; one clitellate (holotype CUMZ 3221) and 10 clitellates (paratypes CUMZ 3222); type locality: river banks of Song River at Vangvieng, Veintiane Province, Laos.

Glyphidrilus bisegmentus Chanabun & Panha, sp. n. (Figs 1, 2A, 3, Table 1) Type material. Holotype: One clitellate (CUMZ 3276) (Fig. 3), Air Banun Pandig, Perak, Malaysia, 496 meters in elevation, S. Panha, R. Chanabun, P. Tongkerd and C. Sutcharit leg., 27 May 2011. 17 clitellates and 8 aclitellates as paratypes (CUMZ 3277), 3 clitellates (ZMH OL14575), 3 clitellates (NHM), same collection data as for holotype. Other material. 22 clitellates and 4 aclitellates (CUMZ 3278), muddy swamp near Kratu waterfall, Kratu, Phuket, Thailand, 07°55'55.5''N, 098°19'23.3"E., 11 m asl, 11 December 2009. Type locality. River banks of Air Banun Pandig, Perak, Malaysia, 05°38'13.4''N, 101°42'41.1"E, 496 m asl. Etymology. This new species was named after the character of having wings only on two segments at 18–19. Diagnosis. Small sized limicolous earthworm with distinct expanded tissues of clitellar wing organs on the lateral sides of the body in 18–19. Clitellum in 16, 17–23, 24. Female pores, male pores and spermathecal pores not visible. Genital markings absent. Four pairs of seminal vesicles in 9–12, with the pair in segment 12 largest. Intestinal origin in 15. Ovaries in 13–14. Prostates, accessory glands, and spermathecae absent. “*” Data same as the holotype “?” Not shown in original description “1” Type specimens examined and data from original description “2” Data only from original description and Shen & Yeo (2005) “3” Data only from original description “4” Type specimen examined Description of holotype. Dimensions: body length 71 mm, diameter 2.3 mm in segment 8, 2.4 mm before clitellar wings in segment 17, 2.0 mm after wings in segment 20 within the clitellum; body cylindrical in anterior part, quadrangular in transverse section behind clitellum. 217 segments. Body colour pale brown with variations from red to pink at adjacent tissues of wing portion in different individuals of newly collected specimens after placement in 30% ethanol for narcotization. At posterior end dorsal surface considerably broader than ventral. Stout wings like lateral ridges in bc extending through segments 18–19 in clitellar region. Prostomium zygolobous. Dorsal pores absent. Clitellum annular in 17–24. Four pairs of setae per segment from 2, setal formula aa:ab:bc:cd:dd = 1.0:0.5:1.1:0.5:1.2 in segment 8. Female pores, male pores and spermathecal pores not visible. Genital markings absent. Septa 4/5–7/8 thicker, 8/9–11/12 thick, from 12/13 to last segment thin. Gizzard small, globular within 8–9. Intestine enlarged from 15. Dorsal blood vessel anterior to 9. Hearts from 9–11. No nephridia distinguishable in first 14 segments. Four pairs of seminal vesicles in 9–12, pair in segment 12 larger than the others but pair in segments 10 and 11 very small. Ovaries in 13–14. Prostate and accessory glands, and spermathecae absent. Variation. Body length 40–71 mm (57.06±20.50), with 152–252 segments. Wings always in 18–19 (covering 2 segments), clitellum in 16, 17–23, 24 (covering 6–9 segments). Genital markings always absent. Habitat. Found on the shore but in proximity to the river water, in the sandy loam topsoil with 65% sand, 35% silt, with pH 7.2, and also under the water at about 5–10 cm in depth. The bank of river was covered with worm casts at Air Banun Pandig, Perak, Malaysia and at Kratu Waterfall, Kratu, Phuket, Thailand. Distribution. The new species is known from the type locality, Air Banun Pandig, Perak, Malaysia and from a muddy swamp near Kratu waterfall, Kratu, Phuket, Thailand. Remarks. This athecal species is quite distinct from others of the genus, which typically have several spermathecae per segment in several segments and wings on more than two segments. Among the species bearing spermathecae, G. malayanus from the Malay Peninsula is most similar to G. bisegmentus sp. n. in the location of the clitellum. However, the other characters are specifically different. Glyphidrilus malayanus has longer wings in 17, ¾18, 18–⅓21, ⅓22, 22 and spermathecae in 14/15–16/17. Glyphidrilus bisegmentus sp. n. differs from the Laotian athecal G. vangviengensis by the following characteristics: G. vangviengensis has longer wings in 24, 25–31, 32, and longer clitellum in 19, 20–35, 36, 37 (Table 1). Material of other species examined for comparison. Glyphidrilus malayanus Michaelsen, 1902: 35: one clitellate (syntype ZMH V5875); type locality: Malay Peninsula, Lubock Paku, Pahang River. Glyphidrilus vangviengensis Panha & Chanabun, 2011: 216, Figs. 1–3; one clitellate (holotype CUMZ 3221) and 10 clitellates (paratypes CUMZ 3222); type locality: river banks of Song River at Vangvieng, Veintiane Province, Laos., Published as part of Chanabun, Ratmanee, Sutcharit, Chirasak, Tongkerd, Piyoros, Tan, Shau-Hwai Aileen & Panha, Somsak, 2012, Three new species of semi-aquatic freshwater earthworms of the genus Glyphidrilus Horst, 1889 from Malaysia (Clitellata: Oligochaeta: Almidae), pp. 120-132 in Zootaxa 3458 on pages 121-124, {"references":["Michaelsen, W. (1922) Oligochaten aus dem Rijks Museum van Natuurlijke Historie zu Leiden. Capita Zoologica, 1, 1 - 68.","Michaelsen, W. (1896) Oligochaeten. Ergebnisse einer zoologischen Forschungsreise in den Notukken und Borneo, im Auftrage der Senckenbergischen naturforschenden Gesellschaft ausgefuhrt von Dr. Willy Kukenthal. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 23, 193 - 243.","Michaelsen, W. (1902) Neue Oligochaeten und neue Fundorte alt-bekannter. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 19, 1 - 54.","Horst, R. (1892) Earthworms from Malay Archipelago. In Max Weber. Zoologische Ergebnisse einer Reise in Niederlandisch Ost Indien, 3, 28 - 83.","Chanabun, R., Bantaowong, U., Sutcharit, C., Tongkerd, P., Inkavilay, K., James, S. W. & Panha, S. (2011) A new species of semi-aquatic freshwater earthworm of the genus Glyphidrilus Horst, 1889 from Laos (Oligochaeta: Almidae). Tropical Natural History, 11, 213 - 222.","Shen, H. P. & Yeo, D. C. J. (2005) Terrestrial earthworms (Oligochaeta) from Singapore. The Raffles Bulletin of Zoology, 53, 13 - 33."]}

Glyphidrilus peninsularis Chanabun & Panha, sp. n. (Figs 1, 2C, 5, Table 1) Type material. Holotype: One clitellate (CUMZ 3279) (Fig. 5), Sungei Bantang, Johor, Malaysia, 99 meters in elevation, S. Panha, R. Chanabun, P. Tongkerd and C. Sutcharit leg., 21 May 2011. 32 clitellates and 12 aclitellates as paratypes (CUMZ 3280), 3 clitellates (ZMH 14576), 3 clitellates (NHM), same collection data as for holotype. Other material. 45 clitellates and 11 aclitellates (CUMZ 3281), Gua Pulai, Kelantan, Malaysia, 04°74'36.3'N, 101°56'31.8"E, 119 m asl, 26 May 2011. 40 clitellates and 24 aclitellates (CUMZ 3282), Kebun Bunka, Penang, Malaysia, 05°26'21.7''N, 100°17'22.5"E, 74 m asl, 19 May 2011. Type locality. River banks of Sungei Bantang, Johor, Malaysia, 02°19'50.5''N, 103°09'45.1"E, 99 m asl. Etymology. This new species was named peninsularis because of its wide distribution throughout the Malay Peninsula. Diagnosis. Small sized limicolous earthworm with distinct expanded tissues of wings in 22, 23–28, ½29, 29. Clitellum in 17, 18–31, 32. Female pores, male pores and spermathecal pores not visible. Genital markings: midventral unpaired in 17, 18, 19, 20, 21 and 31, 32, lateral paired or asymmetrical on bc in 13, 14, 15, 16, 17, 20, 21, 22 and 29, 30. Four pairs of hearts in 8–11. Four pairs of seminal vesicles in 9–12, with the pair in segment 12 largest. Intestinal origin in 14. Ovaries in 13–14. Prostate and accessory glands absent. Four spermathecae in each of 14/15–17/18. Description of holotype. Dimensions: body length 87 mm, diameter 1.1 mm in segment 8, 2.5 mm before the clitellar wings in segment 22, 2.2 mm after wings in segment 30 in clitellar region; body cylindrical in anterior part, quadrangular in transverse section-behind clitellum. 228 segments. Body colour pale brown with variations from red to pink at adjacent tissues of wing portion in different individuals of newly collected specimens. At posterior end dorsal surface considerably broader than the ventral. Clitellar wings on ventro-lateral part of clitellum in 23–29, 2.1 mm long, about 0.5 mm wide on both sides. Prostomium zygolobous. Dorsal pores absent. Clitellum annular in 18–32. Four pairs of setae per segment from 2, setal formula aa:ab:bc:cd:dd = 0.9:0.5:1.0:0.6:1.6 in segment 8. Female pores, male pores and spermathecal pores not visible. Genital markings: paired or asymmetrical on bc in 15, 22, median unpaired on aa in 19 and 31, 32. Septa 5/6–7/8 thicker, 8/9–12/13 thick and 13/14 to the last segment thin. Gizzard small, globular within 7–8. Intestine enlarged from 14. Dorsal blood vessel anterior to 8. Hearts from 8–11. No nephridia distinguishable in the first eleven segments. Four pairs of seminal vesicles in 9–12, with the pair in segment 12 larger than the others. Ovaries in 13–14. Prostate and accessory glands absent. Spermathecae in 14/15–17/18, a pair on each side per segment. Variation. Body length of type and non-type material 49–94 mm (73.79±31.81), with 206–281 segments. Wings in 22, 23–28, ½29, 29 (covering 6–8 segments), with clitellum in 17, 18–31, 32 (covering 14–16 segments). Genital markings: midventral unpaired in 17, 18, 19, 20, 21 and 31, 32, lateral paired or asymmetrical on bc in 13, 14, 15, 16, 17, 20, 21, 22 and 29, 30. Habitat. Found on the shore but in proximity to the river water, in the loamy sand topsoil with 70.83% sand, 23% silt, 4.1% clay, with pH 7.48 from Sungei Bantang, Johor, Malaysia. In the river banks, in the silt topsoil with 93% silt, 7% clay, with pH 8.01 of Gua Pulai, Kelantan, Malaysia. In the sand topsoil 87.50% sand, 8.33% silt, 4.16% clay, with pH 7.61, from Kebun Bunka, Penang, Malaysia and also under the water at about 10–20 cm in depth. Worm casts were abundant along the river banks at all sites. Distribution. The new species is known from the type locality, Sungei Bantang, Johor, river banks of Gua Pulai, Kelantan, and muddy swamp in Kebun Bunka, Penang, Malaysia. Remarks. Glyphidrilus peninsularis sp. n. is different from other species in the following (see also Table 1): G. kukenthali from Sarawak has wings in 18, ½18, 19–24, ½24, longer clitellum in ½13–34, hearts in 7–11, and spermathecae in 14–18. G. malayanus from Malay Peninsula has shorter wings in 17, ¾18, 18–⅓21, ⅓22, 22, shorter clitellum in 15–25, 26, and spermathecae in 14/15–16/17. G. weberi from Java, Sumatra, Celebes, and Flores has longer wings in 23–½32, clitellum in 16–32, and spermathecae in 13/14–17/18. G. quadrangulus from Sumatra has wings in 19, 20–25, and spermathecae in 13/14, 14/15–15/16, 16/17. G. jacobsoni from Sumatra has wings in 21–26, clitellum in 18–30, and spermathecae in 12/13–16/17. G. horsti from Pulau Berhala, Straits of Malacca, has shorter wings in ½23, 23–27, 28, clitellum in 17, 18, ½18, 19–28, 29, 30, ½31, and spermathecae in 14–17. G. gatesi reported from Johor, Malaysia, has wings in 18, ½19, 19–½24, 24, shorter clitellum in ½17, 17, 18–25, ½26, 26, intestinal origin in 18, and spermathecae in 15–17. G. singaporensis reported from Bukit Timah, Singapore, has shorter wings in 21–25, ½26, 26, clitellum in 18, 19–27, ½28, 29, 31, intestinal origin in 15, and spermathecae in 14–17. G. vangviengensis reported from the river banks of Song River at Vangvieng, Laos, has wings in 24, 25–31, 32, longer clitellum in 19, 20–35, 36, 37, and no spermathecae. G. kotatinggi sp. n. from Kota Tinggi waterfall, has shorter wings and clitellum, three pairs of hearts, and spermathecae in 13/14–16/17. G. bisegmentus sp. n. from Air Banun Pandig, Perak, has shorter wings and shorter clitellum, and genital markings and spermathecae afre absent. Material of other species examined for comparison. Glyphidrilus kukenthali Michaelsen, 1896: 195, pl. 13, Fig. 1; type locality: Borneo, Barem River, Sarawak. Glyphidrilus malayanus Michaelsen, 1902: 35; one clitellate (syntype ZMH V 5875); type locality: Malay Peninsula, Lubock Paku, Pahang River. Glyphidrilus weberi Horst, 1889: 77; 3 clitellates (syntype ZMH V 5097); type locality: Java: Buitenzorg, in humid soil, Sumatra: Manindjan and lake of Singkarah, Flores: Kotting, Celebes: Luwu. Annadrilus quadrangulus Horst, 1892: 44; type locality: Lake Danau diatas (near Alahan Pandjang), Sumatra. Glyphidrilus jacobsoni Michaelsen, 1922: 10; one clitellate and 1 aclitellate (syntype ZMH V 9293), type locality: Sumatra, Korintji, Soengai Koembang. Glyphidrilus horsti Stephenson, 1930: 4; type locality: Pulau Berhala, Straits of Malacca. Glyphidrilus gatesi Shen & Yeo, 2005: 16, Fig. 1; one clitellate (holotype ZRC 1974.12.2.51) and 9 clitellates and 2 aclitellates (paratypes ZRC 1974.12.2.52–62); type locality: Sungei Kayu, swamp forest near River Sedili, Johor, Malaysia. Glyphidrilus singaporensis Shen & Yeo, 2005: 18, Fig. 3; one clitellate (holotype ZRC) and 2 clitellates and 9 aclitellates (paratypes ZRC) and 10 clitellates and 9 aclitellates (topotypes CUMZ 3273); type locality: Jungle Fall Valley, Bukit Timah, Singapore. Glyphidrilus vangviengensis Panha & Chanabun, 2011: 216, Figs. 1–3; one clitellate (holotype CUMZ 3221) and 10 clitellates (paratypes CUMZ 3222); type locality: river banks of Song River at Vangvieng, Veintiane Province, Laos., Published as part of Chanabun, Ratmanee, Sutcharit, Chirasak, Tongkerd, Piyoros, Tan, Shau-Hwai Aileen & Panha, Somsak, 2012, Three new species of semi-aquatic freshwater earthworms of the genus Glyphidrilus Horst, 1889 from Malaysia (Clitellata: Oligochaeta: Almidae), pp. 120-132 in Zootaxa 3458 on pages 129-130, {"references":["Michaelsen, W. (1896) Oligochaeten. Ergebnisse einer zoologischen Forschungsreise in den Notukken und Borneo, im Auftrage der Senckenbergischen naturforschenden Gesellschaft ausgefuhrt von Dr. Willy Kukenthal. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 23, 193 - 243.","Michaelsen, W. (1902) Neue Oligochaeten und neue Fundorte alt-bekannter. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 19, 1 - 54.","Horst, R. (1892) Earthworms from Malay Archipelago. In Max Weber. Zoologische Ergebnisse einer Reise in Niederlandisch Ost Indien, 3, 28 - 83.","Michaelsen, W. (1922) Oligochaten aus dem Rijks Museum van Natuurlijke Historie zu Leiden. Capita Zoologica, 1, 1 - 68.","Stephenson, J. (1930) On some Oligochaeta from Berhala Island in the Straits of Malacca. Miscellanea Zoologica Sumatrana, 48, 1 - 5.","Shen, H. P. & Yeo, D. C. J. (2005) Terrestrial earthworms (Oligochaeta) from Singapore. The Raffles Bulletin of Zoology, 53, 13 - 33.","Chanabun, R., Bantaowong, U., Sutcharit, C., Tongkerd, P., Inkavilay, K., James, S. W. & Panha, S. (2011) A new species of semi-aquatic freshwater earthworm of the genus Glyphidrilus Horst, 1889 from Laos (Oligochaeta: Almidae). Tropical Natural History, 11, 213 - 222."]}

Genus Glyphidrilus Horst, 1889 Type species. Glyphidrilus weberi Horst, 1889, by monotypy, Published as part of Jirapatrasilp, Parin, Prasankok, Pongpun, Sutcharit, Chirasak, Chanabun, Ratmanee & Panha, Somsak, 2016, Two new Cambodian semi-aquatic earthworms in the genus Glyphidrilus Horst, 1889 (Oligochaeta, Almidae), based on morphological and molecular data, pp. 543-558 in Zootaxa 4189 (3) on page 548, DOI: 10.11646/zootaxa.4189.3.5, http://zenodo.org/record/166203, {"references":["Horst, R. (1889) Over eene nieuwe soort order de Lumbricinen door Prof. Max Weber uit nedenl. Indie medegebracht."]}

Key to species of Glyphidrilus from Malay Peninsula 1 Gizzard in 7 or ½7–½8 or 7–8........................................................................... 2 - Gizzard in 8 or 8–9.................................................................................... 4 2 Gizzard in 7; wings in 21–26; clitellum in 18–30; spermathecae in 12/13–6/17............................ G. jacobsoni - Gizzard in ½7–½8 or 7–8; wings; clitellum or spermathecae different............................................ 3 3 Intestine from 15; wings in 23–½32; clitellum in 16–32; spermathecae in 13/14–17/18........................ G. weberi - Intestine from 14; wings in 22, 23–28, ½29, 29; clitellum in 17, 18–31, 32; spermathecae in 14/15–17/18..................................................................................................... G. peninsularis sp. n. 4 Gizzard in 8.......................................................................................... 5 - Gizzard in 8–9....................................................................................... 11 5 Genital markings absent; wings in 19, 20–25.................................................... G. quadrangulus - Genital markings different.............................................................................. 6 6 Spermathecae absent; wings in 24, 25–31, 32; clitellum in 19, 20–35, 36, 37......................... G. vangviengensis - Spermathecae present; wings or clitellum different........................................................... 7 7 First hearts in 7; wings in 18, ½18, 19–24, ½24; clitellum in ½13–34; spermathecae in 14–18............... G. kukenthali - First hearts in 8 or 9; wings; clitellum or spermathecae different................................................ 8 8 First hearts in 8; wings in ½23, 23–27, 28; clitellum in 17, 18, ½18, 19–28, 29, 30, ½31; spermathecae in 14–17.... G. horsti - First hearts in 9; wings; clitellum or spermathecae different.................................................... 9 9 Wings in 21–25, ½26, 26; clitellum in 18, 19–27, ½28, 29, 31 and spermathecae in 14–17................ G. singaporensis - Wings from 17 or 18; clitellum or spermathecae different.................................................... 10 10 Intestine from 16; wings in 17, ¾18, 18–¼21, ¼22, 22; clitellum in 15–25, 26; spermathecae in 14/15–16/17... G. malayanus - Intestine from 18; wings in 18, ½19, 19–½24, 24; clitellum in ½17, 17, 18–25, ½26, 26; spermathecae in 15–17.... G. gatesi 11 Wings in 20, 21–¼26, 26; clitellum in 17, 18–28, 29, 30; spermathecae in 13/14–16/17................ G. kotatinggi sp. n. - Wings in 18–19; clitellum in 16, 17–23, 24; spermathecae absent................................ G. bisegmentus sp. n.

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On a collection of land shells made in Borneo by Mr. Everett, with descriptions of supposed new species. Part II. Zotinidae and Helicinidae. Proceedings of the Zoological Society of London, 1891: 22-47.","Gould, A. A., 1843. Description of land mollusks from the province of Tavoy, in British Burmah, collected by Francis Mason. Proceedings of the Boston Society of Natural History, 1: 137-141.","Gude, G. K. & B. B. Woodward, 1921. On Helicella Ferussac. Proceedings of the Malacological Society of London, 14: 174-190.","Gude, G. K., 1903. Classsified list of the helicoild land shell of Asia, part 6. Journal of Malacology, 10: 45-62.","Hausdorf, B., 1995.A preliminary phylogenetics and biogeographic analysis of Dyakiidae (Gastropoda: Stylommatophora) and a biogeographic analysis of other Sundaland taxa. Cladistics, 11: 359-376.","Hemmen, J. & C. Hemmen, 2001 Aktualisierte liste der terrestrischen Gastropoden Thailands. 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