European Journal of Taxonomy 149: 1-34

http://dx.doi.org/10.5852/ejt.2015.149

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2015 • Taylor P.D. & Tan S.H.A.

Research article

urn:lsid:zoobank.org:pub:26669D73-E861-4C8F-99B9-ED286006E853

Cheilostome Bryozoa from Penang and Langkawi, Malaysia

Paul D. TAYLOR 1 & Shau-Hwai Aileen TAN 2

1 Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK.

Corresponding author: [email protected]

2 Marine Science Laboratory, School of Biological Sciences, Universiti Sains Malaysia, 11800 Minden,

Penang, Malaysia. Email: [email protected]

1 urn:lsid:zoobank.org: author: 7AFF2929-DF5B-46B2-94E6-B26B396CC2C8

2 urn:lsid:zoobank.org:author:FB8279A2-D0D7-4151-A30E-81761FA26709

Abstract. Twenty-three species of cheilostome bryozoans are described from the Malaysian islands

of Penang and Langkawi based on a brief reconnaisance survey of shore localities. These are the first

bryozoans to be formally described from either island and they demonstrate the potential for further

research on these neglected suspension feeders. Of the 23 species recorded, 12 are anascans, half of

which are malacostegines, and 11 are ascophorans. The new combinations Acanthodesia falsitemiis

(Liu, 1992), A. perambulata (Louis & Menon, 2009) and A. irregulata (Liu, 1992) are introduced. Most

of the species recorded are widespread in the Indo-Pacific, and some are apparently globally distributed

in the tropics and subtropics, including the invasive fouling species Bugula neritina , Hippoporina indica

and Schizoporella japonica , as well as the coral reef associates Cranosina coronata and Hippopodina

feegeensis. Plastic debris and glass bottles were encrusted by Jellyella eburned, a coloniser of floating

biological and man-made objects that is becoming widespread in the tropics and subtropics of the

world’s oceans.

Keywords. Taxonomy, Bryozoa, Cheilostomata, fouling, Malaysia.

Taylor P.D. & Tan S.H.A. 2015. Cheilostome Bryozoa from Penang and Langkawi, Malaysia. European Journal

of Taxonomy 149: 1-34. http://dx.doi.org/10.5852/eit.2015.149

Introduction

Bryozoans are a phylum of colonial metazoans, numbering almost 6000 species (Bock & Gordon 2013)

and found in all of the world’s oceans. Knowledge of bryozoan faunas is still remarkably poor for some

parts of the world. Among these are the seas around Malaysia. For example, the review of coastal marine

biodiversity in Malaysia by Mazlan et al. (2005) contains no mention of bryozoans. As bryozoans

are common fouling animals, this is clearly anomalous and undoubtedly reflects lack of study rather

than the true absence of these benthic suspension feeders. Bryozoans provide habitats and the food

for numerous other marine animals (e.g., Bradstock & Gordon 1983; Cocito 2004; Wood et al. 2013),

including co mm ercial species, and therefore concern about their vulnerability to enviro nm ental change

is not entirely academic. In particular, most bryozoans have calcareous skeletons that in tropical species

include an apparently high proportion of species biomineralizing aragonite or magnesium-calcite (Taylor

et al. 2015), minerals susceptible to ocean acidification because of their relatively high solubilities.

European Journal of Taxonomy 149: 1-34 (2015)

Penang is an island of 1048 km 2 in the Strait of Malacca; Langkawi, 80 km to the north, is an

archipelago of islands in the Andaman Sea, both off the west coast of the Malaysian Peninsula (Fig. 1).

With one exception, a survey of the literature failed to find any mention of bryozoans from either Penang

or Langkawi; the only reference to bryozoans that could be found was by Waters (1885) in a paper

describing fossil bryozoans from South Australia in which the presence of one species - identified as

Membranipora savartii Audouin, 1826 - is recorded from an unspecified locality in Penang.

Accordingly, a short reconnaisance survey of the shores of Penang and Langkawi was undertaken in

October 2013 to ascertain the presence of bryozoans, identify the species present, and obtain material

for mineralogical analysis to evaluate vulnerability to ocean acidification. The aim of this paper is to

describe the species found, thereby increasing the awareness in Malaysia of these neglected marine

animals and enlarging known species distributions.

A total of 23 cheilostome species is recorded and described from Penang and Langkawi for the first time.

Many of these are invasive fouling species that are known to be widely distributed.

Fig. 1 . Map showing sampling localities in Langkawi (top left) and Penang (bottom left), and the

locations of these islands in Malaysia (right).

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

Material and methods

Localities

Shore collections were made from the following localities in Penang and Langkawi (Fig. 1).

Penang

1. Batu Uban (5°22’ N, 100° 19’ E): a fishing wharf located on the eastern side of the island next to the

main bridge between the peninsula and the island.

2. Batu Maung (5°27’ N, 100° 17’ E): a fishing wharf at the eastern side of the island sheltered by a

peripheral island (Pulau Jerejalc).

3. Pulau Betong (5° 18’ N, 100° IE E): a commercial oyster aquaculture centre on floating cages off the

coast of the small island of Pulau Betong southwest of the main island of Penang.

4. Balik Pulau (5°22’ N, 100° 12’ E): sandy beach in the southwestern part of Penang.

Langkawi

1. Pantai Beringin (6° 17’ N, 99°5E E): sheltered sandy beach in a small bay on the southeastern part of

the island with rocky shore at both ends of the beach.

2. Kuah jetty (bMS’ N, 99°51’ E): small wooden jetty adjacent to the main ferry terminal.

3. Pekan Rabu (6° 19’ N, 99°5CT E): floating dock built from modular plastics connected to each other

and used as a docking area for small local boats.

4. Sungai Menghulu (6° 19’ N, 99°48' E): a fishing wharf at the river mouth leading to the Straits of

Malacca in the south of the island.

5. Kampung Kuala Temoyong (6° 17’ N, 99°44’ E): a fishing wharf at the river mouth leading to the

Straits of Malacca in the southwest of the island.

6. Teluk Bam (6° 15’ N, 99°44’ E): a large wharf located in the southern part of the island used for tourist

and fishing boats.

7. Pantai Chenang (6° 17’ N, 99°43’ E): exposed sandy beach at the southern tip of the island, forming a

narrow shoreline frequented by tourists.

8. Pantai Pasir Hitam (6°25’ N, 99°47’ E): sandy beach with fine sand in the north of the island. A reef

flat composed of discontinuous growing corals is located about 3CM10 m seawards from the beach.

The reef is highly sedimented with fine sand.

Repository

All figured specimens are deposited in the collections of the Marine Science Laboratory, Universiti

Sains Malaysia, Penang (abbreviated MSL). Letters following the number are used to indicate different

species encrusting the same specimen.

Study methods

Collections were made by examining all available artificial and natural hard substrates evident at the

localities visited, using a hand lens to confirm the presence of bryozoans. Only a small proportion

of mollusc shells was found to be bryozoan-encrusted, but artificial substrates, such as pier wharves

and fishing floats, were very commonly fouled by bryozoans. Specimens were washed and dried after

collection. After preliminary study with a binocular microscope, specimens of each species present

were chosen for scanning electron microscopy (SEM). Prior to SEM, colonies were bleached for up to a

day in a dilute solution of commercial bleach to remove soft tissues and expose the skeleton. The exact

time of bleaching was adjusted according to the species concerned and the requirement to conserve

features such as delicate spines while removing cuticular components including the opercula. Bleached

colonies were studied uncoated using a LEO 1455-VP scanning electron microscope and employing a

backscattered electron detector. Measurements were made from SEM images.

European Journal of Taxonomy 149: 1-34 (2015)

Results

Locality records

Penang

Batu Uban

Bugula neritina (Linneaus, 1758)

Batu Maung

Acanthodesia cf. irregulata (Liu, 1992)

Sinoflustra amoyensis (Robertson, 1921)

Pulau Betong

Acanthodesia cf. irregulata (Liu, 1992)

Arbopercula sp.

Parasmittina winstonae Liu et al ., 2001

Parasmittina raigioidea Liu et al. , 2001

Hippoporina indica Pillai, 1978

Scorpiodinipora costulata (Canu & Bassler, 1929)

Balik Pulau

Acanthodesia cf. irregulata (Liu, 1992)

Nellia oculata Busk, 1852

Langkawi

Pantai Beringin

Hippoporina indica Pillai, 1978

Kuah jetty

Cradoscrupocellaria sp.

Celleporaria aperta (Hincks, 1882)

Parasmittina winstonae Liu et al., 2001

Parasmittina sp. 1

Hippoporina indica Pillai, 1978

Schizoporella japonica Ortmann, 1890

Pekan Rabu

Schizoporella japonica Ortmann, 1890

Sungai Menghulu

Hippoporina indica Pillai, 1978

Kampung Kuala Temoyong

Aetea ligulata Busk, 1852

Acanthodesia cf. falsitenuis (Liu, 1992)

Antropora minor (Hincks, 1880)

Adeonella lichenoides (Lamarck, 1816)

Parasmittina sp. 1

Microporella sp.

Scorpiodinipora costulata (Canu & Bassler, 1929)

Teluk Barn

Acanthodesia perambulata (Louis & Menon, 2009)

Hippoporina indica Pillai, 1978

Pantai Chenang

Jellyella eburnea (Hincks, 1891)

Pantai Pasir Hitam

Aetea ligulata Busk, 1852

Acanthodesia perambulata (Louis & Menon, 2009)

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

Cranosina coronata (Hincks, 1881)

Hippopodina feegeensis (Busk, 1884)

Taxonomic descriptions

The descriptions below are based entirely on material collected from Penang and Langkawi and are not

intended to be comprehensive for the species described. Synonymies are abbreviated to include only the

original description plus one key recent reference. All measurements were taken from SEM images of

Malaysian specimens, with ranges of autozooid lengths and widths usually based on 10 zooids.

Order Cheilostomata Busk, 1852

Suborder Inovicellina Jullien, 1888

Family Aeteidae Smitt, 1868

A etea L amouroux, 1812

Aetea ligulata Busk, 1852

Fig. 2A-F

Aetea ligulata Busk, 1852: 31, pi. 42.

Aetea ligulata - Osburn 1950: 13, pi. 1, fig. 4.

Material

MAFAYSIA: MSF BRYOOla, Pantai Pasir Hitam, Fangkawi, collected intertidally from coral reef,

fouling Hippopodina feegeensis. MSF BRY002, Kampung Kuala Temoyong, Fangkawi, encrusting

shell found among fishing debris.

Fig. 2. Aetea ligulata Busk, 1852. A. Colony fouling Hippopodinella feegeensis ; Pantai Pasir Hitam,

Fangkawi, MSF BRYOOla. B-F. Kampung Kuala Temoyong, Langkawi, MSL BRY002. B. Zooids

growing aroung clionid-bored shell. C. Zooid with repaired skeleton. D. Erect part of zooid collapsed

against substrate to reveal annulations on distal side. E. Erect part of zooid collapsed to show opesia on

proximally facing side. F. Broken zooid showing intramural ‘stolon’. Scale bars: A-B = 1 mm; C-E =

200 pm; F = 100 pm.

European Journal of Taxonomy 149: 1-34 (2015)

Description

Colony encrusting, uniserial, runner-like with widely spaced branches (Fig. 2A-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 a nn ulations 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 co mm on cosmopolitan

species Aetea anguina (Linnaeus, 1758) and A sica (Couch, 1844). Amodern 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).

Suborder Malacostegina Levinsen, 1909

Family Membraniporidae Busk, 1852

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.

Acanthodesia cf. falsitenuis (Liu, 1992) comb. nov.

Fig. 3A-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.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

Description

Colony encrusting, multiserial, unilamellar, locally multilamellar. Ancestrula and early astogenetic

stages not observed. Autozooids subhexagonal or subrectangular, distally well-rounded, elongate, 0.53-

Fig. 3. Species of Acanthodesia. — A-C. A. cf. falsitenuis (Liu, 1992). Kampung Kuala Temoyong,

Langlcawi, MSLBRY003. A. Zooids. B. Detail. C. Cryptocystalprocess. —D-F. A.peramabulata (Louis

& Menon, 2009). Pantai Pasir Hitam, Langkawi, MSL BRY004. D. Part of colony with overgrowth.

E. Group of zooids. F. Wide zooid immediately before a row bifurcation. — G-L. A. cf. irregulata (Liu,

1992). Note variation between specimens in the width of the lateral mural rim, which is particularly

broad in I, and the development of tubercles, present only in the colony shown in K and L. G. Pulau

Betong, Penang, MSL BRY005. H-I. Batu Muang, Penang, MSL BRY026. J. Balik Pulau, Penang,

MSL BRY006. K-L. Batu Maung, Penang, MSL BRY007. Scale bars: A, D = 1 mm; B, I, L = 200 pm;

C = 50 pm; E-H, J-K = 500 pm.

European Journal of Taxonomy 149: 1-34 (2015)

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. 3B-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. 3B-C). Similar processes were depicted in zooids of two species from the South

China Sea - Acanthodesia similis (Liu, 1992) (see Liu et al. 2001: pi. 9, fig. 2) and A. falsitenuis (Liu,

1992) (see Liu 1992: fig. 3) - and also in Acanthodesia crenulata (Okada, 1923), described by Tilbrook

(2006: pi. 2b) from the Solomon Islands. Autozooids of A. similis are about half the length of those of

the species described here from Langlcawi, 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.

Acanthodesiaperambulata (Louis & Menon, 2009) comb. nov.

Fig. 3D-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 x 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 comers, 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 encmsting 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.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

Acanthodesia cf. irregulata (Liu, 1992) comb. nov.

Fig. 3G-L

cf. Membranipora irregulata Liu, 1992: 124, figs 14-17.

Biflustra irregulata - Gordon et al. 2007: 46, fig. Id.

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.

3K-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

comers 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. 3K-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. 3K-L) or absent (Fig. 3G-J), and the width of the mural rim varies between specimens

(Fig. 3G-L).

Jellyella Taylor & Monks, 1997

Jellyella eburnea (Hincks, 1891)

Fig. 4

Membranipora eburnea Hincks, 1891: 289, pi. 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 encmsting, multiserial, unilamellar (Fig. 4A, E). Ancestmla 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.

European Journal of Taxonomy 149: 1-34 (2015)

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. 4D-E).

Family Electridae Stach, 1937

Arbopercula Nikulina, 2010

Arbopercula sp.

Fig. 5A-C

Material

MALAYSIA: MSL BRY009, Pulau Betong, Penang, on oyster rafts.

Description

Colony encrusting, multiserial, unilamellar, thinly calcified. Ancestrula and early astogeny not observed.

Autozooids subhexagonal, slender, two or more times longer than wide, 0.49-0.63 mm long by

0.19-0.28 mm wide; boundaries between zooids in adjacent rows marked by a narrow fissure, lacking

between zooids in the same row; opesia occupying nearly all frontal surface, ovoidal; cryptocyst developed

Fig. 4. Jellyella eburnea (Hincks, 1891). Pantai Chenang, Langlcawi. A-C. Scanning electron

micrographs showing variable development of the gymnocyst; MSL BRY008. A. Group of zooids.

B. Slightly oblique view of zooids with poorly developed gymnocysts. C. Zooids with well developed

gymnocysts. D-E. Colonies encrusting a small bottle photographed in the field. Scale bars: A = 1 mm;

B-C = 500 pm.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

only proximally, extremely narrow, steeply sloping; gymnocyst narrow, broadening in proximolateral

corners; two oral spines, erect, short, tapering distally, often with ends closed; circumopesial spines

numbering 6-14, generally paired on either side of opesia, long, tapering distally, overarching frontal

membrane, not overlapping or touching (Fig. 5B); proximal circumopesial spine located close to and

resembling morphologically the oral spines (Fig. 5C).

Remarks

The genus Arbopercula - type species Electra bengalensis Stolizcka, 1869 - was proposed by Nikulina

(2010) for species previously included in Electra , but which have spines on the operculum. These

unmineralized structures have not been observed in the dried material from Penang, which has lost the

opercula. Nevertheless, the Penang bryozoan does show a strong overall resemblance to the specimen

figured by Gordon et al. (2007, fig. 1C) as the type species, particularly in the gymnocystal spinosity, and

also to Arbopercula devinensis (Robertson, 1921), although differences in the distribution and number

of spines, plus the extent of the proximal gymnocyst, preclude unequivocal identification as either of

these two species.

Family Sinoflustridae Gordon, 2009

Sinoflustra Liu & Yang, 1995

Sinoflustra amoyensis (Robertson, 1921)

Fig. 5D-F

Membranipora amoyensis Robertson, 1921: 49, fig. 6.

Sinoflustra amoyensis - Liu & Yang 1995: 349, figs 1-2.

Fig. 5. — A-C. Arbopercula sp. Pulau Betong, Penang, MSL BRY009. A. Group of zooids. B. Zooids

showing spines. C. Distal end of a zooid with a pair of oral spines and the proximal circumopesial

spine of the next distal zooid. — D-F. Sinoflustra amoyensis (Robertson, 1921). Batu Muang, Penang,

MSL BRY010. D. Part of a colony. E. Detail of a group of zooids at the broken edge. F. Opesia and

cryptocystal spinules. Scale bars: A, D = 500 pm; B, E = 200 pm; C, F = 100 pm.

European Journal of Taxonomy 149: 1-34 (2015)

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: pi. 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: pi. 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.

Suborder Flustrina Smitt, 1868

Family Calloporidae Norman, 1903

Cranosina Canu & Bassler, 1933

Cranosina coronata (Hincks, 1881)

Fig. 6

Membranipora coronata 'Hincks, 1881: 147, pi. 10, fig. 1.

Cranosina coronata - Chimonides & Cook 1994: 44, fig. la.

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

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

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.

Family Antroporidae Vigneaux, 1949

Antropova Norman, 1903

Antropova minor (Hincks, 1880)

Fig. 7A

Membranipora trifolium var. minor Hincks, 1880: 87, pi. 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 comer or less often at two proximolateral corners of most autozooids;

rostrum rounded, a little longer but narrower than opesia.

Fig. 6. Cranosina coronata (Hincks, 1881). Pantai Pasir Hitam, Langkawi, MSL BRY011. A. Part of

a colony encrusting a cobble. B. Group of zooids. C. Autozooid with distal avicularium. Scale bars:

A = 1 mm ; B = 500 pm; C = 200 pm.

European Journal of Taxonomy 149: 1-34 (2015)

Remarks

Large vicarious avicularia sometimes develop in this species (e.g., Tilbrook 1998: fig. 2e) but were

not seen in the studied colony from Langlcawi. Antropora minor is widely distributed in the tropics,

occurring in the Caribbean, South Atlantic, Pacific and Indian Oceans, including East Sumbawa,

Malaysia (Tilbrook 1998).

Family Quadricellariidae Gordon, 1984

Nelli a Busk, 1852

Nellia oculata Busk, 1852

Fig. 7B-E

Nellia oculata Busk, 1852: 18, pi 64, fig. 6, pi 65 bis, fig. 5.

Nellia oculata - Winston et al. 2014: 161, fig. 13.

Material

MAFAYSIA: MSF 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

Fig. 7. — A. Antropora minor (Hincks, 1880). Part of a colony encrusting a large gastropod shell;

Kampung Kuala Temoyong, Fangkawi, MSF BRY012a. — B-E. Nellia oculata Busk, 1852. Balik

Pulau, Penang, MSF BRY013. Internodes from a single colony. B, D. Complete internodes. C. Autozooid.

E. Detail of tiny avicularium with proximal pore. Scale bars: A-B, D = 500 pm; C = 100 pm; E = 20 pm.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

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 ajunior 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.

Family Bugulidae Gray, 1848

Bugula Oken, 1815

Bugula neritina (Linnaeus, 1758)

Fig. 8A-B

Sertularia neritina Linnaeus, 1758: 815.

Bugula neritina - Ryland & Hayward 1977: 162, fig. 78.

Fig. 8. — A-B. Bugula neritina (Linnaeus, 1758). Batu Uban, Penang, MSL BRY014. A. Part of a

colony. B. Detail showing four globose ovicells. — C-F. Cradoscrupocellaria sp. Kuah jetty, Langkawi,

MSL BRY015. C. Colony fragment. D. Zooids near branch tip showing spines and scuta. E. Branches

and vibracular seta (bottom). F. Ovicells, spines and scuta. Scale bars: A, C = 1 mm; B, D = 200 pm;

E = 500 pm; F = 100 pm.

European Journal of Taxonomy 149: 1-34 (2015)

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 etal. 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).

Family Candidae d’Orbigny, 1851

Cradoscrupocellaria Vieira et al., 2013

Cradoscrupocellaria sp.

Fig. 8C-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; gymno¬

cyst 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.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

Superfamily Adeonoidea Busk, 1884

Family Adeonidae Busk, 1884

Adeonella Busk, 1884

Adeonella lichenoides (Lamarck, 1816)

Fig. 9A-G

Eschar a l icheno ides L amarck, 1816: 176.

Adeonella lichenoides - Hayward 1988: 126, figs lc, 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. 9F-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.

Fig. 9. Adeonella lichenoides (Lamarck, 1816). A. Field photograph of colony found among fishing

debris at Kampung Kuala Temoyong, Langkawi. B-G. MSL BRY016, fragment of colony shown in

A. B. Bifurcating branch. C. Group of zooids. D. Vicarious avicularium. E. Young zooids near growing

edge before development of the spiramen. F. Zooids with adventitious avicularia and variably developed

calcified bridges isolating the spiramen. G. Older zooids with fully developed spiramens and variously

orientated adventitious avicularia. Scale bars: B = 1 mm; C = 500 pm; D-G =100 pm.

European Journal of Taxonomy 149: 1-34 (2015)

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; cross¬

bar 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).

Superfamily Lepralielloidea Vigneaux, 1949

Family Lepraliellidae Vigneaux, 1949

Celleporaria Lamouroux, 1821

Celleporaria aperta (Hincks, 1882)

Fig. 10A-F

Schizoporella apertaYLincks, 1882: 126, pi. 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.

Fig. 10. Celleporaria aperta (Hincks, 1882). Kuah jetty, Langkawi, MSL BRY017a. A. View of colony

showing growing edge (top left) and older region (bottom right) with frontal buds. B. Developing zooids

at growing edge. C. Region with frontal buds. D. Orifice of non-ovicellate autozooid and steeply inclined

suboral avicularium with dentate distal edge of rostrum. E. Ovicell. F. Two interzooidal avicularia.

Scale bars: A = 1 mm; B = 200 pm; C = 500 pm; D-F = 100 pm.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

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. 10D-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: pi. 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).

Superfamily Smittinoidea Levinsen, 1909

Family Smittinidae Levinsen, 1909

Parasmittina Osburn, 1952

Parasmittina winstonae Liu et al., 2001

Fig. 11A-L

Parasmittina winstonae Liu et al. , 2001: 623, pi. 55, figs 1-7.

Parasmittina winstonae - Tilbrook 2006: 156, pi. 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. 1 ID). 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

European Journal of Taxonomy 149: 1-34 (2015)

0.10 mm wide, large relative to zooid size, lappets developed distolaterally, lyrula narrow, 17-20 pm

high by 25-37 pm wide at top edge, condyles present about one-third distance along orifice, 15 pm high,

medioproximal edge serrated (Fig. 11K); oral spine bases usually numbering two, occasionally one or

Fig. 11. Parasmittina winstonae Liu et al., 2001. — A-I. Kuah jetty, Langkawi, MSL BRY018. A. Small

colony. B. Tiny colony on same substrate. C. Ancestrula of colony shown in B (note that the prism is

from a disaggregated bivalve shell). D. Early stages of another colony showing five budded autozooids

surrounding the ancestrula. E. Group of zooids. F. Giant avicularium with small plectrum-shaped

adventitious avicularium at right proximolateral corner. G. Young autozooids and small adventitious

avicularia. H. Older autozooids and adventitious avicularia. I. Ovicell. — J. MSL BRY019a, same

locality details; oblique view of growing edge with giant bud. — K-L. Pulau Betong, Penang. K. MSL

BRY020a. Slightly oblique view of orifice showing oral spine bases, condyles and lyrula, with plectrum¬

shaped adventitious avicularia distolaterally. L. MSL BRY021a. Old part of colony with numerous

avicularia. Scale bars: A = 1 mm; B, J = 500 pm; C, F-G, I, K = 100 pm; D-E, H, L = 200 pm.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

lacking; ovicell hyperstomial (Fig. 11H-I), broader than long, 0.13-0.16 mm long by 0.18 mm wide,

containing 25M0 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. 1 IF), 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 one-

third 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. 1 IF) 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 aid 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 Tilbroolc (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.

Parasmittina raigioidea Liu et al. , 2001

Fig. 12A-C

Parasmittina raigioidea Liu et al. , 2001: 629, pi. 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 pm

high by 25 pm wide at top edge, condyles present about one-quarter along orifice; oral spine bases

European Journal of Taxonomy 149: 1-34 (2015)

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 pi. 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.

Fig. 12. Parasmittina spp. — A-C. P. raigioidea Liu et al ., 2001. Pulau Betong, Penang, MSL BRY020b.

A. Part of colony. B. Autozooids and avicularia. C. Orifice with lyrula and proximally directed

adventitious avicularium. — D-F. Parasmittina sp. 1. Kuah jetty, Penang, MSL BRY021b. D. Colony

(bottom) becoming overgrown by a colony of P. winstonae (top). E. Ovicellate zooids and proximally

directed vicarious avicularia. F. Detail showing a proximally directed vicarious avicularium (upper left

of centre) with a proximally directed spatulate adventitious avicularium to its lower right and a distally

directed acuminate adventitious avicularium to the lower right of that. — G-I. Parasmittina sp. 2.

Kampung Kuala Temoyong, Langkawi, MSL BRY012b; encrusting a large gastropod shell. G. Part of

colony. H. Group of zooids. I. Autozooids and an adventitious avicularium. Scale bars: A, H = 500 pm;

B, E-F = 200 pm; C, I = 100 pm; D, G = 1 mm

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

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.

Parasmittina sp. 1

Fig. 12D-F

Material

MALAYSIA: MSL BRY021b, Kuah jetty, Langkawi, on fouled rope hanging from jetty (with

P. winstonae).

Description

Colony encrusting, multiserial, unilamellar. Ancestrula and early astogeny not observed. Autozooids

subhexagonal, small, about 0.30-0.32 mm long by 0.25-0.28 mm wide; boundary walls salient with

a fine fissure; frontal shield convex, coarsely pustulose, marginal areolar pores large, no pseudopores;

secondary orifice elliptical, about 0.10-0.12 mm wide, lappets highest proximolaterally, lyrula variable,

typically narrow, 13—18 pm wide at tip; oral spine bases not observed; ovicell hyperstomial, broad,

about 0.12-0.16 mm long by 0.15-0.20 mm wide, ectooecium with about 30 ridged pores, proximal and

distal edges of ovicell overgrown by cryptocystal calcification. Avicularia of three types: (1) adventitious

acuminate avicularia present on some proximal frontal shields, about 0.11 mm long by 0.07 mm wide,

usually distally or distolaterally directed, rostrum an acute triangle, tip raised, rostral shelf shallow,

cross-bar calcified, opesia semicircular; (2) adventitious spatulate avicularia present on some frontal

shields, about 0.14 mm long by 0.05 mm wide, directed proximally or laterally towards autozooid

median line, rostrum rounded with deep shelf, cross-bar calcified, opesia semicircular; (3) vicarious

avicularia large, 0.23-0.38 mm long by 0.13-0.15 mm wide, scattered, directed proximally, rostrum

initially tapering before becoming parallel-sided, tip well rounded, distal edge slightly serrated, rostral

shelf deep, occupying half or more of rostrum length, cross-bar calcified, concave, opesia relatively

small, crescent shaped (Fig. 12F).

Remarks

It has not been possible to identify this species of Parasmittina despite the combination of distinctive

large vicarious avicularia and small adventitious avicularia of spatulate and acuminate types. Pending a

more detailed study and a comprehensive comparison with the myriad of species of Parasmittina , it is

therefore left in open nomenclature as Parasmittina sp. 1.

Parasmittina sp. 2

Fig. 12G-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.

European Journal of Taxonomy 149: 1-34 (2015)

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. 121). 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.

Family Bitectiporidae MacGillivray, 1895

Hippoporina Neviani, 1895

Hippoporina indica Pillai, 1978

Fig. 13A-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.

Fig. 13. Hippoporina indica Pillai, 1978. - A-D. Kuah jetty, Langkawi, MSL BRY017b. A. Group

of zooids. B. Autozooidal orifice and avicularium. C. Autozooids with closed orifices. D. Ovicell. -

E. Pulau Betong, Penang, MSL BRY022; ovicell. - F. Sungai Menghulu, Langkawi, MSL BRY023;

group of ovicellate zooids. Scale bars: A, C, F = 200 pm; B, D-E = 100 pm.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

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. 13D-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.

Superfamily Schizoporelloidea Jullien, 1883

Family Schizoporellidae Jullien, 1883

Schizoporella Hincks, 1877

Schizoporella japonica Ortmann, 1890

Fig. 14A-F

Schizoporella unicornsis var. japonica Ortmann, 1890: 49, pi. 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 x 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. 14E-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.

European Journal of Taxonomy 149: 1-34 (2015)

Remarks

Ryland et al. (2014) comprehensively redescribed Schtoporella 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.

Family Hippopodinidae Levinsen, 1909

Hippopodina Levinsen, 1909

Hippopodina feegeensis (Busk, 1884)

Fig. 15A-C

Lepralia feegeensis Busk, 1884: 144, pi. 22, fig. 9, 9a, 9b.

Hippopodina feegeensis - Til brook 1999: 451, fig. la-h.

Material

MALAYSIA: MSL BRYOOlb, 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

Fig. 14. Schizoporellajaponica Ortmann, 1890. Kuah jetty, Langkawi. — A. Colony photographed in the

field showing colour when alive. — B-D. MSL BRY024. B. Growing edge with giant buds. C. Group

of zooids. D. Orifice and avicularium. — E-F. MSL BRY019b. E. Group of zooids, two ovicellate.

F. Ovicell. Scale bars: B = 1 mm; C, E = 500 pm; D, F = 100 pm.

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

Malaysian material; elsewhere ancestrula comprising a tetrad of zooids (e.g., Tilbrook 1999: fig. lb).

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

Fig. 15. — A-C. Hippopodinafeegeensis (Busk, 1884). Pantai Pasir Hitam, Langkawi, MSL BRY001b.

A. Zooids near growing edge (top). B. Group of autozooids and an ovicell. C. Autozooidal orifice with

an adventitious avicularium. — D-F. Microporella sp. Kampung Kuala Temoyong, Langkawi, MSL

BRY012d. D. Group of zooids. E. Ascopore, orifice and avicularium (top right). F. Oblique view of

ovicellate autozooid showing ascopore and avicularium with mandible. — G. Scorpiodinipora costulata

(Canu & Bassler, 1929) (left) and Microporella (right). MSL BRY012c and MSL BRY012d, respectively.

Kampung Kuala Temoyong, Langkawi. — H-I. Scorpiodinipora costulata (Canu & Bassler, 1929).

H. Kampung Kuala Temoyong, Langkawi, MSL BRY012c. Zooids at the growing edge. I. Pulau Betong,

Penang, MSL BRY025. Group of zooids. Scale bars: A, G = 1 mm; B = 500 pm; C-D, H-I = 200 pm;

E-F = 100 pm.

European Journal of Taxonomy 149: 1-34 (2015)

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.

Family Microporellidae Hincks, 1879

Microporella Hincks, 1879

Microporella sp.

Fig. 15D-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, Caceres-Chamizo & Sanner,

TAYLOR P.D. & TAN S.H.A., Malaysian bryozoans

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.

Superfamily Celleporoidea Johnston, 1838

Family Hippoporidridae Vigneaux, 1949

Scorpiodinipora Balavoine, 1959

Scorpiodinipora costulata (Canu & Bassler, 1929)

Fig. 15G-I

Schizoporella costulata Canu & Bassler, 1929: 317, pi. 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.

Discussion

There appear to be no descriptions in the literature of bryozoans from either Penang or Langkawi,

and the only mention of a bryozoan that could be found from either island is in a paper (Waters 1885:

287) describing Australian fossils in which a bryozoan identified as Membranipora savartii Audouin

(probably one of the species of Acanthodesia described above) is recorded in Penang. The notion that

bryozoans might be rare was dispelled by the discovery of 23 species, some abundant, during just a few

days of shore collecting. Given the economic importance of the marine environment for these islands, it

is remarkable that bryozoans, many of which are significant foulers of man-made structures, have been

totally neglected in the past.

Of the 23 species collected during the current shore survey, 14 were found only at Langkawi, 6 only at

Penang and 3 at both islands. In terms of taxonomy, 12 of the species recorded are anascan cheilostomes

European Journal of Taxonomy 149: 1-34 (2015)

and 11 ascophorans. Half of the anascans are malacostegines, a group characterised uniquely among

living cheilostomes by the possession of planktotrophic, cyphonautes larvae. The high proportion of

malacostegines may be due to the long-lived larvae, allowing colonization of floating substrates by some

species (notably Jellyella eburnea ), and also the ability of other species to tolerate low and fluctuating

salinity environments (e.g., Winston 1977), such as those with high levels of freshwater run-off

Surprisingly, no cyclostome bryozoans were found, but the failure to record soft-bodied ctenostomes

may simply be due to the fact that most samples were collected in a dry condition.

Only 4 of the 23 species have erect colonies, the majority being encrusters. Unique in having robustly

calcified erect colonies is Adeonella lichenoides. This ascophoran was found at one locality on Tangkawi

among what appeared to be fishing by-catch. It seems possible that it came from deeper waters further

offshore than the other bryozoans recorded here. This is supported by the 5.5-113 m total depth range

of the material of A. lichenoides listed by Hayward (1988).

Several of the species collected are known to be invasive foulers. These include Bugula neritina,

Hippoporina indica and Schizoporella japonica, which are becoming more widespread at the present

day (see, e.g., http: //invasions. si. edu/nemesis/browseDB/Group Summary, j sp? G R P=B rvozoans ) . The

impacts of these species on the indigenous biota, as well as on economic fisheries and on ships and

man-made structures such as wharves, are in general poorly understood and completely unknown in

Malaysia.

Appreciation of the true diversity of bryozoans around Penang and Tangkawi and their geographical and

habitat distributions wifi require considerably more research, employing denser sampling and diving and

dredging to obtain bryozoans from subtidal habitats including reefs. For instance, cobbles from subtidal

soft bottoms in nearby southwest Thailand were found by Sanfilippo et al. (2011) to be colonized by a

rich fauna of bryozoans. No comparable sampling for bryozoans has yet been done in Malaysia.

Acknowledgements

We are very grateful for field assistance from students Ivan Chiron Yaman, Woo Sau Pinn and Poi Khoy

Yen. Four reviewers provided comments that led to significant improvements in this paper.

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Manuscript received: 18 May 2015

Manuscript accepted: 7 September 2015

Published on: 3 November 2015

Topic editor: Rudy Jocque

Desk editor: Danny Eibye-Jacobsen

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, De nm ark.