the known and anticipated bryozoan diversity of bangladesh

J. Taxon. Biodiv. Res. 1(2): 45-58, 2007 

THE KNOWN AND ANTICIPATED BRYOZOAN DIVERSITY OF BANGLADESH 

Dennis P. Gordon 1 , Md M. Maruf Hossain 2 , and Tim Wood 3 

1 National Institute of Water & Atmospheric Research, Private Bag 14901 Kilbirnie, Wellington, New Zealand 

2 Institute of Marine Sciences & Fisheries, University of Chittagong, Chittagong 4331, Bangladesh 

3 Department of Biological Sciences, Wright State University, 3640 Colonel Glenn Highway, Dayton, Ohio 45435, USA 

ABSTRACT 

As of early 2007, no member of the phylum Bryozoa had been formally recorded from Bangladesh at the species level, 

either from the 200 nm Exclusive Economic Zone, or from any of the abundant fresh waters. Here we report on 20 taxa 

of Cheilostomata and two of Cyclostomata from the southeastern coast, mostly from beach-cast stones at Cox's Bazar, 

with one species from St Martin's Island. Given the diversity of the phylum in modern seas (c. 6000 species) and the 

known abundance of species in tropical waters including the coasts of northeastern India and northwestern Myanmar 

(Burma), one can predict that the bryozoan fauna of Bangladesh should comprise at least 200 species, and probably 

many more. Brackish-water species are known from the Sundarbans of West Bengal on mangroves and fully marine 

species typically occupy a broad range of hard substrata in coral-reef environments, as well as on seaweeds and 

human debris. Some species are adapted to live on soft sediments. All such forms are expected in the waters of 

Bangladesh. 

Key words: Bryozoa, marine, freshwater, Cheilostomata, Ctenostomata, Phylactolaemata, Cox's Bazar, St. Martin's Island. 

INTRODUCTION 

Bryozoans (moss animals, sea mats, and lace corals) 

are lophotrochozoan protostomes that form colonies made up 

of box-like or tubular modules called zooids. These have 

various forms, but the fundamental unit, a feeding zooid 

(autozooid), comprises a body wall enclosing a polypide 

(lophophore and digestive tract). Bryozoan colonies are mostly 

encrusting, on rocks, shells, and seaweeds, but they can also 

be bushy, resembling hydroids or turfing algae, or large and 

foliaceous like some corals. Some species are free-living, 

forming small discus-like colonies on offshore sand. Deep-sea 

species on soft bottoms have rootlet-like rhizoids that attach to 

individual sediment grains, thereby anchoring the colony in 

the absence of a firm substratum. The largest marine 

bryozoans, like those of Pentapora fascialis (Pallas, 1766) on the 

west coast of Britain and in the Mediterranean, form colonies 

exceeding a metre in diameter (Hayward and Ryland, 1999), 

while the smallest are tiny runner-like colonies with 

microscopic zooids, either on rock or shell or boring into 

shells. Among freshwater species, colonies of Pectinatella 

magnifica in Japan can attain 30 cm across. Most fully grown 

bryozoans fall within a size range of 0.5 cm to a few 

centimeters. Depending upon habitat and substratum, large 

bryozoan colonies can provide habitat for other organisms, 

particularly in temperate waters (e.g. Bradstock and Gordon, 

1983), but in coral reefs bryozoans tend to be dominated by the 

larger scleractinian corals. 

No member of the phylum Bryozoa has been formally 

recorded from Bangladesh at the species level, either from the 

200 nm Exclusive Economic Zone, or from any of the abundant 

freshwaters. Tomascik (1997) reported four genera of 

Cheilostomata from St Martin's Island (Narikel Jinjira) at the 

extreme southeast of Bangladesh. These were recognized from 

field samples to belong to the genera Scrupocellaria (Candidae), 

1 Corresponding author : e-mail: d.gordon@niwa.co.nz 

45 

Stylopoma (Schizoporellidae), and Reteporellina and 

Triphyllozoon (Phidoloporidae) but none was identified to 

species and samples were not kept (Tomascik, pers. comm. 

2007). 

Given the diversity of the phylum in modern seas (c. 

6000 species), especially in tropical to warm-temperate waters, 

it is clear that a significant number of bryozoans can be 

expected in Bangladeshi waters. Similarly for fresh waters - 

although there are fewer species of freshwater bryozoans (c. 85 

worldwide), many of them are fairly widely distributed and 

must certainly exist in the abundant fresh waters of 

Bangladesh. Based on what is known of the bryozoan fauna of 

the coasts of northeastern India and northwestern Myanmar 

(Burma) and the distribution of substrata suitable for 

bryozoans, one can predict that the bryozoan fauna of 

Bangladesh should comprise at least 200 species, and probably 

many more. Brackish-water species are known from West 

Bengal, in tidal ponds and on mangroves in the Sundarbans 

(Annandale, 1907a,b,e; 1908a; Robertson 1921) and fully 

marine species typically occupy a broad range of hard 

substrata in shore environments, as well as seaweeds and 

human debris. Some species are adapted to live on soft 

sediments. All such forms are expected in the waters of 

Bangladesh. 

Recently, a collection of marine bryozoans was made 

from beach-cast sandstone rocks and pumice at Cox's Bazar. 

The small rocks were tunnelled by pholad borings in which a 

number of small bryozoan colonies were concealed. Fracturing 

of the rocks and microscopic examination of the borings 

yielded 21 species, an unexpectedly high diversity for such an 

extensive sandy shoreline. The main purpose of this paper is 

to report on this bryofauna, which represents the first such 

collection made from the coast of Bangladesh. One other 

species collected from St Martin's Island is also reported herein.

D. P. Gordon, M. M. M. Hossain and T. Wood 

MATERIALS AND METHODS 

Selected portions of colonies of beach-collected marine 

bryozoans, or fragments of rock with small colonies, were 

soaked in liquid domestic bleach (commercial sodium 

hypochlorite solution) until organic tissues were removed 

from zooid skeletons (several hours). Colony fragments were 

then washed thoroughly in water and dried prior to carbon 

and Au-Pd coating and scanning electron microscopy. 

Measurements of zooids and other optically resolvable 

features were made using a calibrated eyepiece graticule in a 

binocular microscope and compared with dimensions 

obtained by SEM. 

SYSTEMATIC ACCOUNT OF NEWLY COLLECTED 

MARINE BRYOZOA 

With the exception of Watersipora arcuata, all of the 

species described below were found at Cox's Bazar beach on 

small rocks (sandstone and pumice). 

Phylum Bryozoa Ehrenberg, 1831 

Class Gymnolaemata Allman, 1856. 

Order Cheilostomata Busk, 1852 

Suborder Malacostegina Levinsen, 1902 

Superfamily Membraniporoidea Busk, 1852 

ELECTRIDAE Stach, 1937 

Conopeum ponticum Hayward, 2001 

(Fig. 1A) 

Conopeum ponticum Hayward, 2001:996-997, fig. 1A-D. 

Diagnosis: Colony encrusting, multiserial. Zooids 

subrectangular, arranged quincuncially 0.33-0.69 mm long, 

0.21-0.37 mm wide; opesia elongate-oval, completely 

surrounded by a thin granular cryptocyst that is wider 

proximally and abuts, or nearly abuts, the distal cryptocystal 

rim of the parent zooid; gymnocyst smooth, mostly confined 

to the proximolateral corners. No spines. 

Remarks: This species was first described from Lizard 

Island, Great Barrier Reef, on the basis of cyphonautes larvae 

and small colonies grown from them. The ancestrula is single 

and Hayward (2001) described astogeny as "characteristic of ... 

Conopeum". The species was illustrated only by light 

microscopy. A useful diagnostic feature is the obliteration or 

near obliteration of the midproximal gymnocyst by the 

abutting of adjacent cryptocysts. These characters are shared 

by Aplousina inornamentata Tilbrook, 2006, which appears to be 

conspecific. Conopeum ponticum may turn out to be a 

widespread Indo-Pacific species; colonies have recently been 

collected in northern New Zealand from the hulls of 

recreational vessels plying tropical South Pacific islands 

(Gordon et al. in press). 

Electra bellula (Hincks, 1881) 

(Fig.1B) 

Membranipora bellula Hincks, 1881:149, pl. 8, fig 4. 

Electra bellula: Hastings 1930:706, pl. 2, fig. 8; Liu et al. 2001:437, 

pl. 15, figs 3-6. 

Diagnosis: Colony encrusting, pluriserial. Zooids 

broadly claviform to subpyriform, tapering proximally, 

0.27-0.45 mm long, 0.17-0.29 mm wide. Opesia occupying 

about half the length of the zooid, oval with a steeply sloping 

narrow cryptocyst than can be faintly ribbed laterally. 

Gymnocyst smooth, bearing 3-5 spine bases at the proximal 

end of the opesia, 1-2 spines at the proximal end of the 

gymnocyst, and 0-2 additional spines frontally. 

Remarks: Only the basal encrusting portions of this 

species were found on a beach-cast piece of sandstone in 

pholad borings. Zooids in the present material closely 

resemble a zooid illustrated by Liu et al. (2001, pl. 15, fig. 6) 

from China. The species has been accorded a wide distribution 

from Australia, through the Indo-Pacific, to the Caribbean and 

Brazil and shows considerable morphological variation, not 

only in spination but also in the extent of the proximal 

cryptocyst. It remains to be seen if all of the morphs represent 

the same species. 

46 

Electra bengalensis (Stoliczka, 1869) 

(Fig. 1C) 

Membranipora bengalensis Stoliczka, 1869:55, pl. 12, figs 1-8; 

Thornely 1907:186, fig. 4; Annandale 1907e: 198. 

Electra anomala Osburn, 1950:36, pl. 3, fig. 6. 

Electra bengalensis: Cook 1968:141; McCann et al. 2007:326, fig. 5 

(cum syn.). 


subrectangular, arranged quincuncially, 0.51-0.56 mm long, 

0.14-0.31 mm wide; opesia elongate-oval, bordered by a very 

thin smooth cryptocyst of equal width throughout; gymnocyst 

smooth, mostly confined to the proximolateral corners, very 

narrow or obliterated laterally. Hollow spines around the 

opesial rim (4-5 pairs in the present material) and 1-2 stouter 

spines proximally by the distolateral corners of the orifice of 

the parent zooid. 

Remarks: Only two small colony fragments were found 

of this species, which has up to 11-14 gymnocystal spines in 

well-developed colonies and a diagnostic pair of cervicorn 

spines on the operculum. Originally described from Port 

Canning, West Bengal, this species is known from West Africa, 

Balboa at the Pacific end of the Panama Canal, Florida, 

Queensland and China. 

MEMBRANIPORIDAE Busk, 1852 

Biflustra irregulata (Liu, 1991) 

(Fig. 1D) 

Membranipora irregulata Liu, 1991:57, 78, fig. 1A-E (cum syn.); 

Liu 1992:124, figs 14-18; Liu in Liu et al. 2001:416, pl. 10, 

figs 3 6. 

?Biflustra irregulata: Tilbrook 2006:354. 


rectangular, arranged quincuncially 0.40-0.56 mm long,

Figure 1. A, Conopeum ponticum. B, Electra bellula. C, Electra bengalensis. D, Biflustra irregulata. E, Jellyella 

eburnea. F, Jellyella tuberculata. G, Antropora minor. 

Scale bars: A, C, E-G, 0.25 mm; B, D, 0.20 mm 

47 

Known and anticipated bryozoan diversity of Bangladesh


0.22- 0.36 mm wide; opesia elongate-oval, bordered by a very 

narrow cryptocystal rim laterally and a shelf proximally; 

cryptocyst densely granular-tubercular. No trace of 

gymnocyst. No spines or polymorphs. 

Remarks: A colony was found encrusting Celleporaria 

aperta (Hincks, 1882). According to Liu (1991, 1992), the 

membranous frontal wall and operculum are covered in 

cuticular spinules in life and a pair of smooth tubercles may 

occur on the proximal corners of some zooids (see Liu et al. 

2001, pl. 10, figs 3 and 4). The drawing by Liu (1992, fig. 18) 

and electron micrograph of Liu et al. (2001, pl. 10, fig. 3) of the 

putative ancestrula shows one zooid slightly larger than the 

other, which may explain Tilbrook's (2006) hesitation 

concerning the attribution of this species to Biflustra, to which 

it otherwise conforms. 

Jellyella eburnea (Hincks, 1891) 

(Fig. 1E) 

Membranipora eburnea Hincks, 1891:289, pl. 7, fig. 5. 

Jellyella eburnea: Taylor & Monks 1997: 42, figs 1-13, 16-19 

(cum syn.). 


subrectangular, arranged quincuncially 0.35-0.56 mm long, 

0.22-0.34 mm wide; opesia elongate-oval, partly concealed by 

several stoutly curving gymnocystal tubercles proximally and 

laterally. A smooth narrow cryptocystal shelf occurs under the 

tubercles and around the inner edge of the opesia. No 

articulated spines. 

Remarks: Fresh colonies with intact membranous 

frontal walls occurred on a small pumice rock at Cox's Bazar 

beach. The species is typically pseudoplanktonic on a range of 

floating substrata including seaweeds, Spirula, and Janthina. It 

is widely distributed in the Indo-Pacific from South Africa and 

Kenya to Australasia and is also known from Florida. 

Jellyella tuberculata (Bosc, 1802) 

(Fig. 1F) 

Flustra tuberculata Bosc, 1802:118. 

Membranipora tuberculata: Prenant and Bobin 1966: 115, fig. 29 

(cum syn.). 

Jellyella tuberculata: Taylor and Monks 1997:41, figs 3, 14, 15; 

Tilbrook et al. 2001:37, fig. 2D. 


rectangular to subrectangular, arranged quincuncially, 

0.42-0.52 mm long, 0.19-0.32 mm wide; opesia elongate-oval, 

bordered by a very narrow cryptocystal rim laterally and a 

shelf proximally; cryptocyst granular-tubercular. Gymnocyst 

present proximally, initially at the corners of the zooid, then as 

a thin continuous proximal rim; starting at the corners, the 

gymnocyst arches forward, forming small pockets beneath, 

especially at the corners; in fully calcified zooids the 

gymnocystal tubercles can be stoutly developed, completely 

concealing the proximal cryptocyst. 

48 

Remarks: A single small colony was found at the 

entrance to a broken pholad boring in a small sandstone rock. 

The species is widely distributed in warmer waters of the 

major oceans, often epiplanktonic on Sargassum weed and 

drifting plastic. 

Suborder Neocheilostomatina d'Hondt, 1985 

Infraorder Flustrina Smitt, 1868 

Superfamily Calloporoidea Norman, 1903 

ANTROPORIDAE Vigneaux, 1949 

Antropora minor (Hincks, 1880) 

(Fig. 1G) 

Membranipora trifolium var. minor Hincks, 1880:87, pl. 11, fig. 6. 

Antropora minor: Tilbrook 1998:34, fig. 2 (cum syn.). 

Diagnosis: Colony encrusting, multiserial. Zooids arranged 

more or less quincuncially, 0.28-0.34 mm long, 0.16-0.24 mm wide; 

opesia roundly triangular, c. half the zooidal length; cryptocystal shelf 

extensive, granular-tubercular; gymnocyst restricted to a very thin 

marginal rim and a small proximal tubercle or absent. Small 

interzooidal avicularia found commonly in the angles between three 

zooids, of simple morphology, lacking mandibular pivots or cross-bar. 

Remarks: No ovicells were found. When present, these 

are vestigial and cap-like. The species has a wide distribution 

in the Indo-Pacific and is also known from Brazil. 

Rosseliana sp. 

(Fig. 2A) 


arranged more or less quincuncially, 0.40-0.55 mm long, 

0.16-0.39 mm wide; opesia sunken, subcircular to 

subtrapezoidal, occasionally quite elongate, its border 

rounded distally and straight proximally, occasionally slightly 

constricted in its distal third where the cryptocystal rim curves 

inwardly. Cryptocyst occupying about one third to half the 

zooidal length, its lateral rim curving inwards to divide the 

proximal cryptocyst into two parts, the anterior of which 

slopes inwards to the opesia, the posterior sloping to the 

proximal rim of the zooid. Thin non-articulated spines present, 

each set into the recess at the proximolateral corner of each 

zooid. Avicularia and ovicells not seen. 

Remarks: This is a curious species. It resembles some 

species of Antropora but lacks any trace of a gymnocyst or 

avicularia. It appears closest to the genus Rosseliana Jullien, 

1888, which includes two living species, both 

Atlanto-Mediterranean - the type species R. rosselii (Audouin, 

1826) and R. baeticaensis. Alvarez, 1992), neither of which has 

spines at the zooidal corners, however. In the absence of 

ovicells and other characters, it would be premature to 

describe the present specimen as a new taxon.

Superfamily Microporoidea Gray, 1848 

THALAMOPORELLIDAE Levinsen, 1902 

Dibunostoma reversum (Harmer, 1926) 

(Fig. 2B) 

Calpensia reversa Harmer, 1926:309, pl. 20, figs 13-16 (cum syn.). 

Dibunostoma reversum: Cheetham 1963:52, fig. 27a. 

Dibunostoma reversa: Soule et al. 1991:462, pl. 4, fig. 5; 1992:7. 

Diagnosis: Colony encrusting, multiserial, sheet-like. 

Zooids arranged more or less quincuncially, 0.45-0.62 mm 

long, 0.23-0.36 mm wide; cryptocystal shelf occupying the 

whole frontal area up to the opesia-orifice except for a pair of 

large irregular opesiules, granular-tubercular with c. 25 

pseudopores; gymnocyst restricted to a very thin marginal rim 

and a pair of stout rounded tubercles, one either side of the 

orifice. Interzooidal avicularium sporadic at the bifurcation of 

a zooid row, about one-third the length of an autozooid, 

somewhat square in shape with a small area of cryptocyst, 

short acute rostrum directed proximolaterally, and round to 

oval rostral foramen. No ovicells. 

Remarks: The species has been recorded previously 

only from Sri Lanka, as Thalamoporella rozieri f. falcifera 

(Hincks, 1880) by Thornely (1905). Thalamotreptos Soule, Soule 

& Chaney, 1991 appears to be a junior subjective synonym of 

Dibunostoma Cheetham, 1963. 

Infraorder Ascophorina Levinsen, 1909 

Superfamily Cribrilinoidea Hincks, 1879 

CRIBRILINIDAE Hincks, 1879 

Puellina cf. egretta Ryland & Hayward, 1992 

(Fig. 2C) 

Puellina egretta Ryland & Hayward, 1992: 244, fig. 11d, e; 

Tilbrook et al. 2001: 58, fig. 7C,D; Tilbrook 2006: 94, pl. 

15A, B 

Diagnosis: Colony encrusting, multiserial, the zooids 

somewhat convex with an intervening furrow between 

adjacent zooids. Zooidal length 0.34-0.39 mm, width 0.21-0.27 

mm. Orifice D-shaped, with 5 basally articulated oral spines. 

Frontal shield comprising a spinocyst of 16-18 pinnate costae, 

the adjacent pinnae meeting such that there is a radial series of 

oval foramina between each costal pair; the suboral costae 

bifurcate so as to define a larger subcircular foramen 

resembling an ascopore. Interzooidal avicularia well 

developed, each with an acute rostrum that reaches the second 

oral spine of the adjacent zooid. Ovicells not seen. 

Remarks: Puellina egretta has been accorded a 

widespread Indo-Pacific distribution (Seychelles, Sri Lanka, 

Great Barrier Reef and Coral Sea, Philippines). The present 

material, comprising a single small infertile colony, very 

closely resembles published SEM micrographs of P. egretta but 

has shorter avicularian rostra. More and better-preserved 

material with additional characters is needed from Bangladesh 

for a definite identification. This species will almost certainly 

be found cryptically on coral rubble at St. Martin's Island. 

49 

Known and anticipated bryozoan diversity of Bangladesh 

Superfamily Hippothooidea Busk, 1859 

CHORIZOPORIDAE Vigneaux, 1949 

Chorizopora brongniartii (Audouin, 1826) 

(Fig. 2D) 

Flustra brongniartii Audouin, 1826:240. 

Chorizopora brongniartii: Tilbrook 2006: 102, pl. 17B-D (cum. syn.). 

Diagnosis: Colony encrusting, multiserial, the zooids 

slightly separated from each other by a narrow network of 

tiny connecting tubes and small kenozooids. Zooids 

transparent, arranged more or less quincuncially, 0.38-0.44 

mm long, 0.22-0.28 mm wide. Gymnocystal frontal shield with 

faint transverse striae and usually 1-2 tubercle in the midline. 

Orifice transversely D-shaped. Ovicell helmet-like, 

surmounted apically by a small avicularium with mandibular 

pivots, the ovicell opening closed by the zooidal operculum. 

Interzooidal kenozooids with a circular frontal foramen. 

Remarks: This species has been accorded a 

near-cosmopolitan distribution within temperate and tropical 

waters. 

TRYPOSTEGIDAE Gordon, Tilbrook & Winston in Winston, 2005 

Trypostega henrychaneyi Tilbrook, 2006 

(Fig. 2E) 

Trypostega henrychaneyi Tilbrook, 2006:110, pl. 19D-F (cum syn.). 


hyaline to opaque, diamond-shaped, such that they are widest 

mid-length and tapering distally and proximally, 0.30-0.47 mm 

long, 0.15-0.27 mm wide. Orifice cleithridiate, the median 

sinus relatively long, a narrow rounded V in shape. 

Gymnocystal frontal shield evenly perforated by tiny pores. 

Each zooid with a small middistal zooeciule with a small 

central foramen. Ovicell somewhat flattened, with a distal 

foramen and porous like the frontal shield, the orifice closed 

by the zooidal operculum. 

Remarks: This species is one of the most robust species 

of Trypostega, forming relatively large colonies that can be 

multilaminar. Previously confused with European T. venusta 

(Norman, 1864), Tilbrook (2006) has shown that it has a 

relatively wide Indo-Pacific distribution from the Red Sea 

through Indonesia and Queensland to Fiji, eastern Polynesia 

and Panama. 

Superfamily Lepralielloidea Vigneaux, 1949 

LEPRALIELLIDAE Vigneaux, 1949 

Celleporaria aperta (Hincks, 1882) 

(Figure 2, F-G) 

Schizoporella aperta Hincks, 1882:126, pl. 5, fig. 3. 

Celleporaria aperta: Harmer 1957:673 (part); Winston and 

Heimberg 1986:32, figs 79-84; Ryland & Hayward 

1992:251, fig. 14d,e.


Figure 2. A, Rosseliana sp. B, Dibunostoma reversum. C, Puellina cf. egretta. D, Chorizopora brongniartii. E, Trypostega 

henrychaneyi. F, G, Celleporaria aperta. 

Scale bars: A, D, E, 0.25 mm; B, 0.30 mm; C, F, G, 0.10 mm. 

50

Diagnosis: Colony forming self-overgrowing 

multilamellar crusts of somewhat irregularly orientated 

zooids, these c. 0.39-0.47 mm long, 0.19-0.22 mm wide. Frontal 

shield smoothly textured with small tubercles; circular 

marginal areolar pores present around the periphery of the 

shield. Orifice with a distinct short rounded sinus, its outer 

corners somewhat cusp-like, the anter with a pair of small but 

distinct condyles; the sinus typically partly concealed by an 

umbo in which there is a small transversely orientated 

avicularium with complete crossbar, the rostrum rounded, 

toothed. A pair of oral spine bases present, one at each 

distolateral corner of the orifice. Ovicell hood-like, less 

tubercular than the frontal shield. Vicarious avicularia with a 

moderately elongate tapering rostrum, the tip rounded, a 

complete crossbar present. 

Remarks: Celleporaria aperta is widespread in the 

tropical and subtropical Indo-pacific 

Superfamily Smittinoidea Levinsen, 1909 

SMITTINIDAE Levinsen, 1909 

Pleurocodonellina signata (Waters, 1889) 

(Fig. 3A) 

Smittia signata Waters, 1889:17, pl. 3, figs 4-6. 

Pleurocodonellina signata: Tilbrook 2006:170, pl. 36C-F (cum syn.). 

Diagnosis: Colony encrusting. Zooids 0.33-0.50 mm 

long, 0.20-0.34 mm wide, the umbonuloid frontal shield 

coarsely dimpled frontally, bordered by 13-19 areolar pores, 

initially round to subelongate and becoming larger with 

secondary calcification. Orifice with a somewhat irregular, 

raised peristomial rim in older zooids, with very stout 

inwardly projecting, minutely denticulate condyles that define 

a drop-shaped sinus. Oral spine bases three in marginal 

zooids. Adventitious avicularia single or paired, subjacent to 

the orifice on one or both sides and directed proximally, of 

variable length, typically narrow with a rounded rostral tip 

and complete crossbar; palatal shelf half the length or more of 

the rostrum. Ovicell recumbent, with the endooecium 

exposed, with numerous pores, each set in an excavation, the 

rim often slightly raised; typically an irregular wide band of 

secondary calcification across the proximal face of the ovicell 

distal to the orificial rim which is slightly raised above the 

level of the ovicell. 

Remarks: Tilbrook (2006) has illustrated the holotype 

specimen of P. signata, to which the present material very 

closely conforms. 

Parasmittina sp. 

(Fig. 3B) 

Diagnosis: Colony encrusting, the zooids subquadrate 

to elongate and tapering, 0.29-0.36 mm long, 0.18-0.21 mm 

wide. Frontal shield umbonuloid, bordered by 15-18 

conspicuous marginal areolar pores. Orifice with a relatively 

narrow lyrula. Oral spine bases not evident. Relatively large 

51 


adventitious avicularium present subjacent to one corner of 

the orifice. Ovicells not seen. 

Remarks: Only one small worn colony was found. But 

for apparently larger avicularian chambers, it resembles P. 

galerita Ryland and Hayward, 1992, known from Queensland 

and Vanuatu, but its condition and size preclude certain 

identification. One consistent feature in the present colony is 

the occurrence of an areolar pore, pertaining to the distal 

zooid, on the mid-distal rim of the orifice, the raised 

peristomial rim of which abuts the areolar pore on each side at 

a higher level. 

WATERSIPORIDAE Vigneaux, 1949 

Watersipora arcuata Banta, 1969 

Watersipora arcuata Banta, 1969:96, figs 1-4 (cum syn.); Gordon 

1989:41, pl. 20J (cum syn.). 

Diagnosis: Colony encrusting, black in life, with 

vermilion growing edge. Zooids more or less 

elongate-rectangular, 0.68-1.11 mm long, 0.29-0.60 mm wide, 

the frontal shield evenly perforated by numerous 

pseudopores, with a slightly larger pore on each side near the 

proximal corner of the orifice. Orifice slightly wider than long. 

No oral spines, avicularia or ovicells. 

Remarks: This well-known warm-water species was 

photographed at St Martin's Island by M. Hossain. First 

described from San Diego Bay, California, the species is 

known from the Galapágos, New Zealand, New South Wales, 

and Queensland (Gordon and Mawatari 1992). 

Superfamily Schizoporelloidea Jullien, 1883 

MICROPORELLIDAE Hincks, 1879 

Microporella harmeri Hayward, 1988 

(Fig. 3C) 

Microporella harmeri Hayward, 1988: 324, pl. 10e. 

Diagnosis: Colony encrusting, the zooids 0.38-0.49 mm 

long, 0.25-0.49 mm wide, generally widest at midlength. 

Frontal shield granular-tubercular, with small, simple 

pseudopores scattered between the granulations. Orifice 

transversely D-shaped, with a minutely denticulate proximal 

rim and 3-4 small spine bases distally. Ascopore crescentic 

with tiny spinules. Avicularia single, placed subjacent to the 

ascopore, the width variable, with a truncated spout-like 

rostral tip and complete crossbar. Ovicell skeletal surface like 

frontal shield, with numerous tiny pseudopores; a smooth 

band of calcification along its proximal rim above the orifice. 

Remarks: Microporella harmeri was first described from 

Mauritius. Its most distinctive features are the denticulate 

proximal rim of the orifice in combination with a proximally 

broad avicularian cystid. Hayward (1988) showed the mandible


(not seen in the present material) as being unusually short and 

hastate. He did not depict the ovicell, however, which is 

illustrated here for the first time. 

Superfamily Celleporoidea Johnston, 1838 

HIPPOPORIDRIDAE Vigneaux, 1949 

Odontoporella sp. 

(Fig. 3D) 

Diagnosis: Colony encrusting, the zooids 0.24-0.31 mm 

long, 0.14-0.20 mm wide, widest in its proximal third. Frontal 

shield relatively smooth and imperforate frontally, bordered 

by c. 6 small areolar pores marginally. Orifice relatively large, 

longer than wide, with a pair of small condyles not quite 

mid-length, separating the anter from the broad rounded 

poster. No oral spines or ovicells. Avicularia not seen. 

Remarks: Carter and Gordon (2007) have recently 

reviewed the austral genus Odontoporella, concluding that 

there are three species - from Chile, New Zealand, and from 

New Caledonia. The present material, comprising a single tiny 

colony fragment, resembles New Zealand O. bishopi Carter 

and Gordon but is unlikely to be this species. More and better 

material is needed before the Bangladesh species can be 

properly characterised. It represents the first record of the 

genus from the Northern Hemisphere. 

CELLEPORIDAE Johnston, 1838 

Turbicellepora sp. 

(Fig. 3E-F) 

Diagnosis: Colony encrusting, the zooids recumbent at 

the colony margin, becoming more erect in the centre of the 

colony, 0.38-0.56 mm long, 0.28-0.39 mm wide. Frontal shield 

smooth, imperforate except for marginal areolar pores, these 

more evident in the distal half. Orifice longer than wide, with 

a rounded broadly V-shaped sinus, set within a continuous 

thin peristomial rim. No oral spines. Avicularia per se not 

evident; a pair of kenozooidal structures present in the place 

of avicularia, one on either side of the peristome; occasionally 

just a single larger such structure on one side. Ovicell sunken, 

the broad endooecial tabula perforated by more than 30 small 

pores. 

Remarks: The present material almost certainly 

represents a new species but the two small incomplete colony 

fragments (both less than 5 mm maximum length) are 

inadequate for erecting a new taxon. 

In discussing a new species of Turbicellepora from the 

Solomon Islands, Tilbrook (2006) remarked that the presence 

of the genus in the tropical Indo-Pacific is questionable, 

suggesting that it might be appropriate including such species 

in another genus. One such genus potentially exists - 

Scorpiodinipora Balavoine, 1959 (type species Cellepora bernardii 

Audouin, 1826), but as Hayward (1988) pointed out, this 

genus may be synonymised with Turbicellepora (regardless of 

52 

attributed misidentification on the part of Balavoine, though 

this is not certain - see also d'Hondt 2006). Not withstanding, 

existing Indo-Pacific Turbicellepora species conform to their 

Atlanto-Mediterranean congeners in all essential features. On 

the other hand, the present material from Cox's Bazar lends 

support to Tilbrook's suggestion - it has the ovicellular pores 

confined to a tabular area, which is not the case for all other 

species of Turbicellepora. It is beyond the scope of the present 

paper to comment further on generic status without recourse 

to a revision of the genus. 

PHIDOLOPORIDAE Gabb and Horn, 1862 

Rhynchozoon attina Tilbrook, 2006 

(Fig. 4A-C) 

Diagnosis: Colony encrusting. Zooids 0.41-0.35 mm 

long, 0.21-0.25 mm wide, the frontal skeletal surface 

porcellanous but irregular owing to sparse tubercles; c. 5 pairs 

of marginal areolar pores present. Orifice with a beaded distal 

rim, the proximal sinus a rounded V shape, relatively broad, 

the condyles stout, shoulder-like, continuous with the 

beading. No articulated oral spines, even at the colony margin. 

Peristomial rim asymmetrical, with a pseudosinus on one side 

and bearing three projections. A suboral avicularium set 

transversely into the peristome, visible frontally; one or a pair 

of acute avicularia on the frontal shield, generally at its widest 

point, orientated transversely, with thin crossbar. Ovicells 

immersed, usually completely concealed by secondary 

calcification, with a vertically descending labellum. 

Remarks: Rhynchozoon attina was first described from 

the Solomon Islands. It superficially resembles a species of 

Exochella Jullien, owing to the frequently paired mid-lateral 

avicularia that are directed laterally. The most distinctive 

features of the species are the orientation of the avicularia, the 

peristomial pseudosinus in ephebic zooids, and the shape of 

the orifice. The present material may have stout frontal 

tubercles not described by Tilbrook in his specimens. In this 

regard, it resembles the species from Komodo attributed to R. 

verruculatum (Smitt, 1873) by Winston and Heimberg (1986), 

but Smitt's species is from the Caribbean and Smitt depicted a 

deeper sinus. The material from Komodo is here regarded as 

belonging to Tilbrook's species. 

Class Stenolaemata Borg, 1926 

Order Cyclostomata Busk, 1852 

Suborder Tubuliporina Milne Edwards, 1838 

Superfamily Tubuliporoidea Johnston, 1838 

DIASTOPORIDAE Gregory, 1899 

Plagioecia sp. 

(Fig. 4D-E) 

Diagnosis: Colony encrusting, radial. Zooids with 

tubular peristomes, width ~0.07-0.08 mm, some of them with 

perforated terminal diaphragms, the perforations with tiny 

spinules. Zooidal surface punctuated by pseudopores, each 

with thin radii. Partial, broken brood chamber present, more

Figure 3. A, Pleurocodonellina signata. B, Parasmittina sp. C, Microporella harmeri. D, Odontoporella sp. 

E, F, Turbicellepora sp. 

Scale bars: A-D, 0.15 mm; E, 0.25 mm; F, 0.2 mm. 

53 

Known and anticipated bryozoan diversity of Bangladesh


Figure 4. A-C, Rhynchozoon attina note the orificial sinus appears somewhat shallower than in frontal view 

owing to tilting; in C, ov indicates the prsence of ovicells. D, E, Plagioecia sp.; E showing peristomial 

diaphragms. F, indeterminate rectangulate cyclostome. 

Scale bars: A, C, E, 0.15 mm; B, 0.02 mm; D, 1.00 mm; F, 0.4 mm. 

54

densely porous than autozooids. 

Remarks: The present material is very worn and long 

peristomes are lacking. The lack of a brood chamber and other 

morphological details precludes further identification, but the 

terminal diaphragms are typical of Plagioecia. 

Suborder Rectangulina Waters, 1887 

Superfamily Lichenoporoidea Smitt, 1867 

INCERTAE SEDIS 

Genus et species indeterminate 

(Fig. 4F) 

Diagnosis: Colony encrusting, radiate, with a narrow 

flattened band of differentiating zooids around the periphery. 

Autozooids in narrow uniserial radiating rows separated by 

larger-diameter alveolae chambers, their walls becoming 

thickened in the colony interior; zooidal peristomial width 

~0.06-0.08. Both zooidal peristomes and alveoli have pinhead 

spinules on the interior walls. 

Remarks: In the absence of a brood chamber further 

determination is not possible. Pinhead spinules are known in 

several rectangulate taxa, including species of Disporella Gray 

(Lichenoporidae) and also in Favosipora holdsworthii (Busk, 

1875) (Densiporidae), which is known from Sri Lanka (Alvarez 

1992). 

DISCUSSION 

Anticipated bryozoan diversity of Bangladesh 

Bryozoan literature (Annandale 1906, 1907a-e, 1908a-c, 

1909a,b, 1910, 1911a,b, 1912, 1915; Thornely 1907; Annandale 

and Kemp 1912; Robertson 1921; Ganapati and Satyanarayana 

Rao 1968; Satyanarayana Rao and Lakshmana Rao 1969; 

Satyanarayana Rao and Ganapathi 1972a,b, 1975, 1978, 1980, 

1986; Satyanarayana Rao et al. 1984l; Satyanarayana Rao and 

Balaji 1988; Satyanarayana Rao 1991, 1992; Samanta 1999) has 

been consulted to gaguge known bryozoan diversity in the 

waters immediately south of Bangladesh, based on historical 

collections from the coast of northeastern India (West Bengal 

and Orissa states and Andhra Pradesh north of 

Vishakhapatnam) and northwestern Burma, and for brackish 

and fresh waters. These areas have not been adequately 

sampled and, after corrections for synonyms and ignoring taxa 

of equivocal status (e.g. species that were not illustrated or 

poorly so, and doubtful biogeographical records), only about 

44 marine bryozoans so far known from the Indian and 

Burmese sectors of the northern Bay of Bengal can also be 

expected to occur in the 166,000 km 2 Bangladesh EEZ. 

The muddy to sandy continental shelf of Bangladesh 

(with an area of 66,400 km 2 , of which 36 % is = 10 m depth) has 

not been explored bryozoologically, though free-living or 

rooted forms may be anticipated. Rare hard-bottom outcrops 

at shelf break or in deeper water to beyond 1000 m should also 

yield bryozoans. A handful of specially adapted estuarine 

bryozoans are expected for the tidally active Bangladesh 

55 


Sundarbans. Some bryozoans have also been recorded from 

marine turtles (e.g. Lepidochelys olivacea (Eschscholtz) - Frazier 

et al. 1992) and the xiphosuran Tachypleus gigas (Patil and Anil 

2000) in the Bay of Bengal. 

It has become appreciated in recent years that 

bryozoan diversity can be very high in coral environments 

(Winston 1986; Ryland and Hayward 1992; Hayward and 

Ryland 1995; Tilbrook 2006). Apart from some turfing, 

robustly encrusting, or prominent erect species, however, 

most bryozoans are small and cryptic, living on the undersides 

of coral heads or worn coral boulders, often in the spaces 

between branches or in excavations provided by other 

organisms. On inner reef flats, conditions are unsuitable for 

bryozoans, but on the outer flat where boulders and rubble are 

common, bryozoans can be found in abundance, especially at 

the reef edge at low water and on the seaward slope (Jackson 

et al. 1985; Winston 1986; Ryland and Hayward 1992). Where 

space allows, as on the underside of concave surfaces, 

relatively large, erect colonies can grow. Their major roles on 

reefs are as hidden encrusters, cavity dwellers, cavity fillers, 

and dead-coral veneerers; between reefs they can act as 

sediment binders, stabilisers, and trappers (Ryland 1974; 

Cuffey 1977). Hence, they can contribute in a small way to reef 

build-up, especially those that engage in self-overgrowth or 

which grow over other organisms. Such colonies tend to 

become a permanent part of the calcareous mass that 

comprises a reef, acting as accessory frame-builders. 

Alternatively, those bryozoans that break up upon the death 

of the colony can contribute to the sediment that forms 

between clumps and reefs, just as they do in more temperate 

settings (Wass et al. 1970). It is anticipated that most of the 

undiscovered bryozoan diversity for Bangladesh will be found 

in the vicinity of St. Martin's Island (Tomascik, 1997). 

Freshwater bryozoans have not yet been reported in 

the literature from Bangladesh, but at least eight species can be 

anticipated based on their occurrence in waters that pass 

through Bangladesh or which have been reported in other 

countries in the wider region. Certainly, a number of species 

must occur in the abundant small lakes, ponds, and reservoirs 

in Bangladesh. Based on a museum specimen from Rajshahi, 

only one species record (Plumatella casmiana) exists for 

Bangladesh, reported here for the first time. Annandale (1912) 

reported P. casmiana Oka as P. testudinicola new species, on 

carapaces of Gangetic cheloninas at Rajmahal, India, just 

upstream of the border with Bangladesh. Co-occurring with 

P. casmiana on the crowned river turtle Hardella thurjii (Gray) 

was the ctenostome bryozoan Hislopia lacustris Carter, which 

can certainly be expected within the borders of Bangladesh. 

REFERENCES 

Allman, G. J. 1856. A Monograph of the Freshwater Polyzoa, 

Including all the Known Species, both British and foreign. 

The Royal Society, London. viii + 119pp., 11 pls. 

Alvarez, J. A. 1992a. Briozoos de la Campaña Fauna I. Parte 

primera : Ctenostomida y Cheilostomida Anascina. 

Cah. Biol. Mar. 33 : 273-297.


Alvarez, J. A. 1992b. Sobre algunas especies de la familia 

Lichenoporidae Smitt, 1866 (Bryozoa, Cyclostomida) en 

el región Atlántico-Mediterránea. Parte I : género 

Disporella Gray, 1848. Cah. Biol. Mar. , 33: 201-243. 

Annandale, N. 1906. Notes on the freshwater fauna of India. 

No. 11. The affinities of Hislopia. J. Asiatic Soc. Bengal, 2: 

59-63. 

Annandale, N. 1907a. Notes on the freshwater fauna of India. 

No. 12. The Polyzoa occurring in Indian fresh and 

brackish pools. J. Asiatic Soc. Bengal, 3: 83-93, pl. 2. 

Annandale, N. 1907b. The fauna of brackish ponds at Port 

Canning, Lower Bengal. Part I.-Introduction and 

preliminary account of the fauna. Rec. Ind. Mus., 1: 

35-43. 

Annandale, N. 1907c. Further note on a polyzoon from the 

Himalayas. Rec. Ind. Mus., 1: 145-148. 

Annandale, N. 1907d. Statoblasts from the surface of a 

Himalayan pond. Rec. Ind. Mus., 1: 177. 

Annandale, N. 1907e. The fauna of brackish ponds at Port 

Canning, Lower Bengal. Part VI.-Observations on the 

Polyzoa, with further notes on the ponds. Rec. Ind. Mus. 

, 1: 197-205. 

Annandale, N. 1908a. The fauna of brackish ponds at Port 

Canning, Lower Bengal. Part VII.-Further observations 

on the Polyzoa, with the description of a new genus of 

Entoprocta. Rec. Ind. Mus., 2: 11-19. 

Annandale, N. 1908b. Corrections as to the identity of Indian 

Phylactolaemata. Rec. Ind. Mus., 2: 110. 

Annandale, N. 1908c. Three Indian Phylactolaemata. Rec. Ind. 

Mus., 2: 169-174. 

Annandale, N. 1909a. Preliminary note on a new genus of 

phylactolaematous Polyzoa. Rec. Ind. Mus. , 3: 279-280. 

Annandale, N. 1909b. A new species of Fredericella from Indian 

lakes. Rec. Ind. Mus. 3: 373-374. 

Annandale, N. 1910. Materials for a revision of the 

phylactolaematous Polyzoa of India. Rec. Ind. Mus., 5: 

37-57. 

Annandale, N. 1911a. Freshwater sponges, hydroids and 

Polyzoa. Fauna of British India, 1911: 1-251, 5 pls. 

Annandale, N. 1911b. Systematic notes on the ctenostomatous 

Polyzoa of fresh water. Rec. Ind. Mus., 6: 193-201, pl. 13. 

Annandale, N. 1912. Fauna symbiotic Indica. No. 3.-Polyzoa 

associated with certain Gangetic tortoises. Rec. Ind. 

Mus., 7: 147-150, pl. 13. 

Annandale, N. 1915. The genus Australella and some allied 

species of phylactolaematous Polyzoa. Rec. Ind. Mus., 

11: 163-169, pls 2, 3. 

Annandale, N. 1922. The marine element in the fauna of the 

Ganges. Bijdr. Dierk. Amsterdam 22: 150. 

Annandale, N. and S. Kemp. 1912. Observations on the 

invertebrate fauna of the Kumaon Lakes, with special 

reference to the sponges and Polyzoa. Rec. Ind. Mus., 7: 

129-145. 

Audouin, J. V. 1826. Explication sommaire des planches de 

Polypes de l'Égypte et de la Syrie, publiées par 

Jules-César Savigny [Planches 1-14 par J.-C. Savigny 

(1817)]. Pp. 225-244. In: Description de l'Égypte, ou receuil 

des obsrvations et des recherches qui ont été faites en Égypte 

pendant l'Expedition de l'Armée française, publiée par les 

56 

ordres de sa majesté l'Empéreur Napoléon le Grand. Histoire 

Naturelle. Vol. 1, part 4. Imprimerie Impériale, Paris. 

339 p. 

Balavoine, P. 1959. Bryozoaires. Mission Robert Ph. Dollfuss 

Égypt, Résult. scient., 3(34): 257-282, pls 1-6. 

Banta, W. C. 1969. Watersipora arcuata, a new species in the 

subovoidea-cucullata-nigra complex (Bryozoa, 

Cheilostomata). Bull. So. Calif. Acad. Sci., 68: 96-102. 

Borg, F. 1926. Studies on Recent cyclostomatous Bryozoa. Zool. 

Bidr. Upps., 10: 181-507, 14 pls. 

Bosc, L. A. G. 1802. Histoire naturelle des vers. Vol. 3, 

pp.1-270. In: Buffon, G. L. L. de: Histoire naturelle de 

Buffon, classée ... après le système de Linné ... par R. R. 

Castel ... nouvelle édition. Déterville, Paris. 

Bradstock, M. and D. P. Gordon. 1983. Coral-like bryozoan 

growths in Tasman Bay, and their protection to 

conserve commercial fish stocks. N. Z. J. Mar. Freshwat. 

Res. , 17: 159-163. 

Busk, G. 1852. An account of the Polyzoa, and sertularian 

zoophytes, collected in the voyage of the Rattlesnake, 

on the coasts of Australia and the Louisiade 

Archipelago, &c. Pp. 343-402, pl. 1 (Appendix No. IV). 

In: MacGillivray, J. (ed.), Narrative of the Voyage of 

H.M.S. Rattlesnake, commanded by the late Captain Owen 

Stanley ... 1846-1850; including discoveries and surveys in 

New Guinea, the Louisiade Archipelago, etc., to which is 

added the account of Mr E. B. Kennedy's expedition for the 

exploration of the Cape York Peninsula [including Mr W. 

Carron's narrative]. Vol. 1. T. W. Boone, London. 

Busk, G. 1859. A Monograph of the Fossil Polyzoa of the Crag. The 

Palaeontographical Society, , London. 136pp. 

Busk, G. 1875. Catalogue of the Cyclostomatous Polyzoa in the 

Collection of the British Museum. Trustees of the British 

Museum, London. 41 p., 34 pls. 

Carter, M. C. and D. P.Gordon. 2007. Substratum and 

morphometric relationships in the bryozoan genus 

Odontoporella, with a description of a new 

paguridean-symbiont species from New Zealand. Zool. 

Sci., 24: 47-56. 

Cheetham, A. H. 1963. Late Eocene zoogeography of the 

eastern Gulf Coast region. Geol. Soc. Amer. Mem., 91: 

i-xii, 1-113, 3 pls. 

Cook, P. L. 1968. Bryozoa (Polyzoa) from the coast of tropical 

West Africa. Atlantide Rep., 10: 115-262, pls 8-11. 

Cuffey, R. J. 1977. Bryozoan contributions to reefs and 

bioherms through geologic time. Pp. 181-194. In: Reefs 

and Related Carbonates - ecology and Sedimentology. Frost, 

S. H., Weiss, M. P. and J. B. Saunders (eds). [AAPG 

Studies in Geology No. 4.] The American Association 

of Petroleum Geologists, Tulsa, Oklahoma. 421pp. 

Ehrenberg, C. G. 1831. Symbolae physicae, seu icones et 

descriptiones corporum naturalium novorum aut minus 

cognitorum, quae ex itineribus per Libyam, Aegyptum, 

Nubiam, Dongalam, Syriam, Arabiam et Habissiniam ... 

studio annis 1820-25 redierunt ... Pars Zoologica. Vol. 4, 

Animalia Evertebrata exclusis Insectis. Berlin. 10 pls. 

Frazier, J. G., J. E. Winston and C. A. Ruckdeschel. 1992. 

Epizoan communities on marine turtles. III. Bryozoa. 

Bull. Mar. Sci., 51: 1-8.

Gabb, W. M. and G. H. Horn. 1862. The fossil Polyzoa of the 

Secondary and Tertiary formations of North America. 

J. Acad. Nat. Sci. Philadelph., 5: 111-179, pls 19-21. 

Ganapati, P. N. and K. Satyanarayana Rao. 1968. Fouling 

bryozoans in the Vishakapatnam harbour. Curr. Sci., 

37: 81-83. 

Gordon, D. P. 1989. The marine fauna of New Zealand: 

Bryozoa: Gymnolaemata (Cheilostomida Ascophorina) 

from the western South Island continental shelf and 

slope. Mem. N.Z. Oceanogr. Inst., 97: 1-158. 

Gordon, D. P., A. M. Hosie and M. C. Carter. (In press.) 

Post-2000 detection of warm-water alien bryozoan 

species in New Zealand - the significance of 

recreational vessels. Virginia Mus. Nat. Hist. Mem. 

Gordon, D. P. and S. F. Mawatari. 1992. Atlas of 

marine-fouling Bryozoa of New Zealand ports and 

harbours. Misc. Publs. N.Z. Oceanogr. Inst. 107: 1-52. 

Gray, J. E. 1848. List of the Specimens of British Animals in the 

Collection of the British Museum. Part 1. Centroniae or 

Radiated Animals. Trustees of the British Museum 

(Natural History), London. xiii + 173pp. 

Gregory, J. W. 1899. Catalogue of the Fossil Bryozoa in the 

Department of Geology, British Museum (Natural History). 

The Ctreaceous Bryozoa. Vol. 1. Trustees of the British 

Museum, London. 457pp. 

Harmer, S. F. 1926. The Polyzoa of the Siboga Expedition. Part 2. 

Cheilostomata Anasca. Siboga Exped. 28b: viii, 181-501, 

pls 13-34. 

Hastings, A. B. 1930. Cheilostomatous Polyzoa from the 

vicinity of the Panama Canal collected by Dr. C. 

Crossland on the cruise of the S.Y. 'St. George'. Proc. 

Zool. Soc. Lond., 47: 697-740, pls 1-17. 

Hayward, P. J. 1988. Mauritian cheilostome Bryozoa. J. Zool. 

Lond., 215: 269-356. 

Hayward, P. J. 2001. The identity of two species of 

cyphonautes larvae (Bryozoa: Cheilostomata) from 

Lizard Island, Great Barrier Reef, Australia. J. Mar. 

Biol. Ass. U.K., 81: 995-999. 

Hayward, P. J. and J. S. Ryland. 1995. Bryozoa from Heron 

Island, Great Barrier Reef. 2. Mem. Qld Mus., 38: 

533-573. 

Hayward, P. J. and J. S. Ryland. 1999. Cheilostomatous Bryozoa. 

Part 2. Hippothooidea - Celleporoidea. Synopses Br. 

Fauna, n.s. 14 (2nd edn): 1-416. 

Hincks, T. 1879. On the classification of the British Polyzoa. 

Ann. Mag. Nat. Hist., ser. 5, 3: 153-164. 

Hincks, T. 1880. Contributions towards a general history of the 

marine Polyzoa. II. Foreign Membraniporina. Ann. 

Mag. Nat. Hist., ser. 5, 6: 81-92, 376-384, pls 10-11, 16-17. 


marine Polyzoa. IV. Foreign Membraniporina (second 

series). Ann. Mag. Nat. Hist., ser. 5, 7: 147-161, pls 8-10. 


marine Polyzoa. XV. South-African and other Polyzoa. 

Ann. Mag. Nat. Hist., ser. 6, 7: 285-298, pls 6-7. 


marine Polyzoa. IX. Foreign Cheilostomata 

(miscellaneous). Ann. Mag. Nat. Hist., ser. 5, 9: 116-127, 

pl. 5. 

57 


Hondt, J.-L. d'. 1985. Contribution à la systématique des 

Bryozoaires eurystomes. Apports récents et nouvelles 

propositions. Ann. Sci. nat., Zool. sér. 13, 7: 1-12. 

Hondt, J.-L. d'. 2006. Nouvelles Explications des Planches de 

"Polypes" de la Description de 'Égypte. II. Bryozoaires 

Bryozoaires, Planches 6 à 13. Institut d'Orient, Paris. 

86pp. 

Jackson, J. B. C., J. E. Winston, and A. G. Coates. 1985. Niche 

breadth, geographic range, and extinction of Caribbean 

reef-associated cheilostome Bryozoa and Scleractinia. 

Pp. 151-158. In: Proceedings of the Fifth International 

Coral Reef Congress, (Gabrie, C. and B. Salvai, eds.) 

Antenne Musum-ÉPHÉ, Moorea. 

Johnston, G. 1838. A History of the British Zoophytes. W. H. 

Lizars, Edinburgh, London & Dublin. xii + 341 pp., 44 

pls. 

Jullien, J. 1883. Bryozoaires. Espèces draguées dans l'océan 

Atlantique en 1881. Bull. Soc. Zool. Fr., 7: 497-529, pls 

13-17. 

Levinsen, G. M. R. 1902. Studies on Bryozoa. Vidensk. Medd. 

naturh. For.Kjøbenhavn, 54: 1-31. 

Liu, X. 1991. A study on the anascan-cribrimorphan bryozoans 

from Nansha Islands of China. Bull. Mar. Biol. Nansha 

Is. Neighbouring Seas 1: 56-81. [In Chinese with English 

summary.] 

Liu, X. 1992. On the genus Membranipora (Anasca: 

Cheilostomata: Bryozoa) from south Chinese seas. 

Raffles Bull. Zool., 40: 103-144. 

Liu, X., X. Yin and J. Ma. 2001. Biology of Marine-Fouling 

Bryozoans in the Coastal Waters of China. Science Press, 

Beijing. xxi + 860 p., 82 pls. [In Chinese.] 

Marcus, E. 1937. Bryozoarios marinhos Brasileiros. I. Bol. 

Fac.Phil. Sci. Letr. Univ. São Paulo. Zoologia ,1: 1-162, 

221-224, pls 1-29. 

McCann, L. D., N. G. Hitchcock, J. E. Winston and G. M. Ruiz. 

2007. Non-native bryozoans in coastal embayments of 

the southern United States: new records for the 

western Atlantic. Bull. Mar. Sci. , 80: 319-342. 

Milne Edwards, H. 1838. Mémoire sue les Crisis, les Hornères, 

et plusieurs autres polypes vivants ou fosiles dont 

l'organisation est analogue à celle des Tubulipores. 

Ann. Sci. Nat., Paris 9: 193-238. 

Norman, A. M. 1903. Notes on the natural history of East 

Finmark. Polyzoa. Ann. Mag. Nat. Hist., ser. 6, 13: 

112-133, pls 6-7. 

Osburn, R. C. 1950. Bryozoa of the Pacific Coast of America 

part 1, Cheilostomata-Anasca. Allan Hancock Pacif. 

Exped., 14:1-269. 

Pallas, P. 1766. Elenchus zoophytorum sistens generum 

adumbrationes generaliores et specierum cognitarum 

succinctas descriptiones cum selectis auctorus synonymis. 

Petrum van Cleef, hagae-Comitum. xviii + 451pp. 

Patil, J. S. and A. C. Anil. 2000. Epibiotic community of the 

horseshoe crab Tachpleus gigas. Mar. Biol. , 136: 699-713. 

Prenant, M. and G. Bobin. 1966. Bryozoaires. Faune Fr., 

68(2):1-647. 

Robertson, A. 1921. Report on a collection of Bryozoa from the 

Bay of Bengal and other eastern seas. Rec. Ind. Mus., 

22: 33-65


Ryland, J. S. 1974. Bryozoa in the Great Barrier Reef Province. 

Pp. 341-348. In: Proceedings of the Second International 

Coral Reef Symposium, 1. Cameron, (A. M. et al., eds.) 

Great Barrier Reef Committee, Brisbane. 

Ryland, J. S. and P. J. Hayward. 1992. Bryozoa from Heron 

Island, Great Barrier Reef. Mem. Qld Mus., 32: 223-301. 

Samanta, T. K. 1999. Fresh water Bryozoa. Fauna West Bengal 

10: 445-461. 

Satyanarayana Rao, K. 1991. Marine Bryozoa. Pp. 535-541. In: 

Zoological Survey of India. Animal Resources of India. 

Protozoa to Mammalia: State of the Art, Zoological 

Survey of India, Calcutta. 694pp. 

Satyanarayana Rao, K. 1992. Breeding and seasonal abundance 

of inter-tidal Bryozoa (Ectoprocta) at Visakhapatnam, 

Bay of Bengal. Proc. Natnl. Acad. Sci. Ind., 62B: 535-541. 

Satyanarayana Rao, K. and M. Balaji. 1988. Biological fouling 

at Port Kakinada, Godavari Estuary, India. Pp. 551-574. 

In: Marine Biodeterioration. Advanced Techniques 

Applicable to the Indian Ocean. (Thompson, M.-F., R. 

Saraojini & R. Nagabhushanam, eds.) Oxford & IBH 

Publishing Co., Ltd., New Delhi. xxxii + 826pp. 

Satyanarayana Rao, K. and P. N. Ganapathi. 1972a. Some new 

and interesting bicellariellids (Polyzoa : 

Cheilostomata) from Vishakapatnam coast, Bay of 

Bengal. Proc. Ind. Natl Sci. Acad., 38B(3&4): 21-29. 

Satyanarayana Rao, K. and P. N. Ganapathi. 1972b. On the 

common anascan genus Electra from Vishakapatnam 

and its vicinities. Proc. Ind. Natl Sci. Acad., 38B(3&4): 

220-224. 

Satyanarayana Rao, K. and P. N. Ganapati. 1975. Littoral 

Bryozoa in the Godavari Estuary. Bull. Dept. Mar. Sci., 

Univ. Cochin, 7: 591-600. 

Satyanarayana Rao, K. and P. N. Ganapati. 1978. Biology of 

fouling bryooans at Vishakpatnam harbour. Proc. Ind. 

Acad. Sci., 87B: 63-75. 

Satyanarayana Rao, K. and P. N. Ganapati. 1980. Epizoic fauna 

of Thalamoporella gothica var. indica and Pherusella 

tubulosa (Bryozoa). Bull. Mar. Sci., 30: 34-44. 

Satyanarayana Rao, K. and P. N. Ganapati. 1986. Distribution 

pattern and abundance of Bryozoa on rocky shores at 

Visakhapatnam, Bay of Bengal, India. Ind. Edn Ser., 12: 

563-578. 

Satyanarayana Rao, K. and M. V. Lakshmana Rao. 1969. 

Record of Kinetoskias sp. (Bicellariellidae : Polyzoa) 

from Vishakapatnam coast, Bay of Bengal. Curr. Sci., 

38: 387-388. 

Satyanarayana Rao, K., B. Viswanadham and G. Samuel Raju. 

1984. Some newly recognised intertidal Bryozoa 

(Ectoprocta) from Vishakapatnam, Bay of Bengal. 

Geobios New Rep. , 3: 106-110. 

Smitt, F. A. 1868. Bryozoa marina in regionibus arcticis et 

borealibus viventia recensuit. Ofvers. Kongl. 

Vetenskaps-Akad. Forh. 24: 443-497, pls 16-20. 

Smitt, F. A. 1873. Floridan Bryozoa, collected by Count L. F. de 

Pourtales. Part II. Kongl. Svensk. Vetensk.-Akad. Handl. 

11(4): 1-83, pls 1-13. 

Soule, D. F., J. D. Soule and H. W. Chaney. 1991. Some little 

known genera of Thalamoporellidae: Thairopora, 

58 

Diploporella, and new genera Marsupioporella and 

Thalamotreptos. In: Bigey, F. P. (ed.) Bryozoaires Actuels 

et Fossiles : Bryozoa Living and Fossil. Bull. Soc. Sci. 

Nat. Ouest Fr., Mém. HS, 1: 447-464. 

Soule, D. F., J. D. Soule and H. W. Chaney. 1992. The genus 

Thalamoporella worldwide (Bryozoa, Anasca): 

morphology, evolution and speciation. , 1: 1-92. 

Stach, L. W. 1937. Reports of the McCoy Society for Field 

Investigation and Research. Lady Julia Percy Island. 

13. Bryozoa. Proc. R. Soc. Vict., n.s. , 49: 373-384, pl. 22. 

Stoliczka, F. 1869. On the anatomy of Sagartia Scilleriana and 

Membranipora bengalensis. J. Asiatic Soc. Bengal, 38:28-63. 

Taylor, P. D. and N. Monks. 1997. A new cheilostome 

bryozoan genus pseudoplanktonic on molluscs and 

algae. Invert, Biol. , 116: 39-51. 

Thornely, L. R. 1905. Report on the Polyzoa collected by 

Professor Herdman, at Ceylon, in 1902. Rep. Gov. 

Ceylon Pearl Oyster Fish. Gulf Manaar, 4 (Suppl. Rep. 

26): 107-130, pl. 1. 

Thornely, L. R. 1907. Report on the marine Polyzoa in the 

collection of the Indian Museum. Rec. Ind. Mus., 1: 

179-196. 

Tilbrook, K. J. 1998. The species of Antropora Norman, 1903 

(Bryozoa: Cheilostomatida), with the description of a 

new genus in the Calloporoidea. Rec. S. Austr. Mus., 31: 

25-49. 

Tilbrook, K. J. 2006. Cheilostomatous Bryozoa from the 

solomon Islands. Santa Barbara Mus. Nat. Hist. Monogr., 

(Stud. Biodiv. No. 3) 4: 1-385. 

Tilbrook, K.J.; Hayward, P.J. and Gordon, D.P. 2001. 

Cheilostomatous Bryozoa from Vanuatu. Zool. J. Linn. 

Soc. Lond., 131: 35-109. 

Tomascik, T. 1997. National Conservation Strategy 

Implementation Project - 1. Management Plan for Coral 

Resources of Narikel Jinjira (St. Martin's Island). Final 

Report (Draft for Consideration). Ministry of 

Environment and Forest, Government of Bangladesh, 

Dhaka. 126pp. 

Vigneaux, M. 1949. Révision des Bryozoaires néogènes du 

Bassin d'Aquitaine et essai de classification. Mém. Soc. 

géol. Fr., n.s., 28: 1-153, 11 pls. 

Wass, R. E., Conolly, J. R. and R. J. MacIntyre. 1970. Bryozoan 

carbonate sand continuous along southern Australia. 

Mar. Geol. ,9: 63-73. 

Waters, A. W. 1887. On Tertiary cyclostomatous Bryozoa from 

New Zealand. Q. Jl Geol. Soc. Lond., 43:337-350, pls 6-8. 

Waters, A. W. 1889. Bryozoa from New South Wales. Part IV. 

Ann. Mag. Nat. Hist., ser. 6, 4: 1-24, pls 1-3. 

Winston, J. E. 2005. Re-description and revision of Smitt's 

"Floridan Bryozoa" in the collection of the Museum of 

Comparative Zoology, Harvard University. Virginia 

Mus. Nat. Hist. Mem. 7: i-ix, 1-147. 

Winston, J. E. 1986. An annotated checklist of coral-associated 

bryozoans. Amer. Mus. Novit. No. 2859: 1-39. 

Winston, J. E. and B. F. Heimberg. 1986. Bryozoans from Bali, 

Lombok, and Komodo. Amer. Mus. Novit. No. 2847: 

1-106. 

Manuscript received 02 October 2007 

Rivised version accepted 20 October 2007

the known and anticipated bryozoan diversity of bangladesh

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