Family LANCEOPORIDAE Harmer, 1957
Compiler and date details
May 2015, updated - ABRS
July 2001 - Dr Philip Bock
- Lanceoporidae Harmer, 1957.
Introduction
The family Parmulariidae was introduced for the genus Parmularia alone, by Canu & Bassler (1927). The close relationship of this genus with the genera Emballotheca and Calyptotheca, had been partially discussed by Livingstone (1928), whose conclusions were rejected by Harmer (1957). Emballotheca and Calyptotheca have since been referred to the Schizoporellidae and the Hippoporinidae, but the similarities among the three genera were regarded as close enough by Chimonides & Cook (1993), to include them all together within the family Parmulariidae.
The genus Lanceopora d'Orbigny (1852) is certainly closely allied to Parmularia, but its type species (L. elegans) was based on an immature colony, which is in the collection of the Musee National de Histoire Naturelle (D.P.Gordon, pers. comm.). The question of the synonymy of Parmularia with Lanceopora requires a detailed comparative study, using additional material of the type species of Lanceopora. The decision to replace the family name Parmulariidae with the new name Lanceoporidae, made by Harmer (1957), on the assumption that the two genera were subjective synonyms, appears premature, and has been unnecessary since 1961 (ICZN Art.40).
Parmularia has an unique and distinctive colony form, unlike those found in either Calyptotheca or Emballotheca, which are usually encrusting. However, in Calyptotheca, several species develop as lunulitiform colonies. There are strong similarities among the three genera in such characteristics as range of orifice shape, dimorphism of brooding zooids, frontal shield calcification and ovicell structure. In addition, both Parmularia and Calyptotheca have species which develop vicarious avicularia, some capable of feeding.
The frontal shield is cryptocystidean, with marginal frontal septulae and numerous frontal pseudopores. Secondary calcification may change the appearance of zooids later in ontogeny. The primary orifice may be sinuate, or the proximal edge may be curved distally. There are paired condyles but no oral spines. Ovicells are usually large, are closed by the operculum, and have a porous shield, derived from one or more distal autozooids. Suture lines mark the limits of each of the distal zooid's contribution. The orifices of brooding zooids are often dimorphic and enlarged. Avicularia may be adventitious in Emballotheca and Calyptotheca, and vicarious or interzooidal in both Calyptotheca and Parmularia.
The family has an overwhelmingly Indo-West Pacific distribution. Species assigned to Emballotheca or Calyptotheca from the eastern Pacific do not seem to belong to either of these genera. Species from the Antarctic (Rogick 1955) have been discussed by Hayward & Thorpe (1988) and reassigned to a new genus of Smittinidae, Thrypticocirrus.
Parmularia is principally known from the East Indies and Australia, and is associated with 'sand faunas', where the substratum consists of fine particles only. Colonies of all species are bilaminar, and leaf-like or discoid, up to 4 cms in diameter. The arrangement of zooids on either face is identical, and there is a series of marginal kenozooids with uncalcified frontal shields round the edge of each colony. On the proximal edge, the kenozooids become extrazooidal, and some extrazooidal calcification develops as colonies grow larger. The colony is anchored within the upper few cms of the surface sediment, by a large, cuticle-covered, turgid, extrazooidal rhizoid structure which may be 2-3 mms wide and up to 10 cms long. The major part of the rhizoid is buried in the sediment, and it ends in terminal fibrils which adhere to the sand grains etc. In some large colonies, additional, secondary rhizoids develop. The rhizoid is capable of repairing damage to its cuticle, and of regaining its turgidity, by which it supports the bilaminar head of the colony vertically above the surface of the sediments (Bock 1982), and Wass (1984). Ovicells are regularly derived from 1-3 distal zooids, and often show sutures. The orifices of the brooding zooids are usually wider than those of autozooids.
No adventitious avicularia occur in Parmularia, and not all species develop large vicarious avicularia. In both P. quadlingi and P. occidenta, these avicularia are elongated and rounded distally, with paired peg-like condyles ( Chimonides & Cook 1993).
The most commonly distributed species is P. obliqua (Bock 1982, as Lanceopora). Eschara reniformis Kirchenpauer 1869 may be a synonym, but MacGillivray's description was published earlier (Harmer 1957: 984). The species has no vicarious avicularia and is reported from northwestern and Western Australia to Bass Strait and Tasmania. Bock (1982) noted that it is 'common in water deeper than about 20m'. The growth and mortality of P. obliqua on mobile sand 'megaripples' in Spencer Gulf, South Australia, was investigated by Shepherd (1983). The species was identified as an r-strategist with high rates of recruitment, mortality and growth. P. obliqua has been illustrated in colour by Bock (1982) and Wass (1984).
Four species have been reported from the Queensland coast. Of these, P. arnoldi differs from all others in having autozooid orifices elongated like the avicularia of P. quadlingi and the secondary thickening of the frontal shield obscures the frontal pores, and large mucrones are developed.
P. arnoldi has been observed alive, and the settlement of larvae and their early development of colonies described. The large orange colonies develop large numbers or ovicells late in astogeny. The larvae begin searching behaviour among the sand grain substratum within 15-30 minutes of release. After a maximum of 30 more minutes they burrow actively into the sand. Larvae adhere to a sand grain, and metamorphose within 12 hours into a minute rhizoid and a trilobed head of primary autozooid buds. Young colonies five days after release develop a 3 mm rhizoid and a bilaminar head of 8 zooids (Cook & Chimonides 1985).
The large entangled mass of rhizoid fibrils and sand grains anchoring fully grown colonies houses numerous colonies, at all astogenetic states, of small rooted species of Sphaeropora and Conescharellina. All these forms are thus capable of a completely interstitial life.
The genus Calyptotheca was introduced by Harmer (1957) for Schizoporella nivea var. wasinensis Waters (1913) from East Africa. The characters defining this generic group had already been discussed by Rogick (1955) and both Calyptotheca and Emballotheca were further revised and described by Dumont (1981). Calyptotheca is characterised by the sinuate primary orifice. In all other characters it is closely similar to Emballotheca.
A large number of species described from Australia appear to referable to Calyptotheca but require revision. At present 5 species are known from South Australia and Victoria. These include C. variolosa, first described as Cellepora by MacGillivray (1869), and later, as Schizoporella biturrita by Hincks (1884). This species was illustrated by Bock (1982) as Gigantopora biturrita (Gordon & Parker 1991). C. variolosa encrusts 'stems and fronds of algae in shallow water'. C. anceps and C. subimmersa (MacGillivray, 1879) and C. triangula (Hincks, 1881) are also known from southern Australia. The species illustrated by Wass & Yoo (1983) as Schizomavella lata is actually C. triangula. They gave a distribution extending from Western Australia to New South Wales. However, a species called Schizoporella lata by MacGillivray (1883) from Victoria appears to belong to Calyptotheca. At least another six species are known from the Queensland coast. These include C. hastingsae Harmer (1957), from Torres Strait and C. australis (Haswell 1881), which was referred to Schizomavella by Hastings (1932), also from the Northern Territory and the Great Barrier Reef. Ryland & Hayward (1992) and Hayward & Ryland (1995){RYLANDJ1995A} illustrated three further species from Heron Island, at the south end of the reef: C. tenuata Harmer (1957), C. thornelyae Dumont (1981) and C. rupicola. The species described as Metroperiella pyriformis by Harmer (1957) from Holborne Island and Torres Strait also appears to be referable to Calyptotheca. Finally, a free-living, lunulitiform species was described from northwest Australia by Cook (1965) as C. conica. C. conica has autozooids with adventitious avicularia; large vicarious avicularia, with viscera and tentacles are also present.
The genus Emballotheca was introduced by Levinsen (1909) for Eschara quadrata MacGillivray (1880) from Victoria. Emballotheca has almost the same suite of characters as Calyptotheca, except that the proximal side of the primary orifice is curved distally. Adventitious avicularia are usually present, but vicarious avicularia have not been found in any species. E. quadrata had originally been described by MacGillivray in 1869, but under a preoccupied name, Eschara elegans. E. quadrata was figured by Wass & Yoo (1983) from the western Bass Strait. Rogick (1955) noted that Mucronella quadrata Busk (1884) was not the same species, and renamed it Emballotheca buskii. This name predates the name E. pacifica used in the same way by Harmer (1957); see Dumont (1981). E. buskii Rogick (1955) was also figured by Ryland & Hayward (1992) (as E. pacifica), from Heron Island. Both E. quadrata and E. buskii have wide Australian distributions.
The Parmulariidae seems to have closest similarities with genera included in the Bitectiporidae, viz. Metroperiella, and the Gigantoporidae viz. Gigantopora.
Emballotheca occurs from the Victorian Tertiary (MacGillivray 1895; Canu & Bassler 1935), and Maplestone (1910) described fossil specimens of Parmularia from the same area.
Diagnosis
Colony erect, moderately well calcified, either supported and anchored by an extrazooidal, turgid, cuticular peduncle and additional rhizoids, or encrusting, erect and tubular. Zooids either in bilaminar, round or leaflike lobes, with marginal kenozooids, or unilaminar. Zooid frontals lepralioid (cryptocystidean), with numerous pseudopores and sinuate orifice. Avicularia adventitious and vicarious, sometimes with tentacles and viscera. Brooding in large ovicells derived from several distal zooids, with sutures and pores frontally.
General References
Bock, P.E. 1982. Bryozoans (Phylum Bryozoa). pp. 319-394 in Shepherd, S.A. & Thomas, I.M. (eds). Marine Invertebrates of Southern Australia. Handbook of the Flora and Fauna of South Australia Adelaide : Government Printer Part 1 491 pp.
Busk, G. 1884. Polyzoa. Pt. I. Cheilostomata. Report on the Scientific Results of the Voyage of H.M.S. Challenger 1873–1876, Zoology 10: xiv, 216
Canu, F. & Bassler, R.S. 1927. Classification of the cheilostomatous Bryozoa. Proceedings of the United States National Museum 69(14): 1-42
Canu, F. & Bassler, R.S. 1935. New species of Tertiary cheilostome Bryozoa from Victoria, Australia. Smithsonian Miscellaneous Collections 93(9): 1-54
Chimonides, P.J. & Cook, P.L. 1993. Notes on Parmularia MacGillivray (Bryozoa: Cheilostomata) from Australia. Memoirs of the Queensland Museum 33: 41-48
Cook, P.L. 1965. Notes on some Polyzoa with conical zoaria. Cahiers de Biologie Marine 6: 435-454
Cook, P.L. & Chimonides, P.J. 1985. Larval settlement and early astogeny of Parmularia (Cheilostomata). pp. 71-78 in Nielsen, C. & Larwood, G.P. (eds). Bryozoa: Ordovician to Recent. Fredensborg : Olsen & Olsen.
Dumont, J.P.C. 1981. A report on the cheilostome Bryozoa of the Sudanese Red Sea. Journal of Natural History 15: 623-637
Gordon, D.P., & Parker, S.A. 1991. Discovery and identity of 110-year-old Hutton Collection of South Australian Bryozoa. Records of the South Australian Museum (Adelaide) 25: 121-128
Harmer, S.F. 1957. The Polyzoa of the Siboga Expedition. Part 4. Cheilostomata Ascophora II. Siboga-Expéditie Report 28D: 641-1147
Hastings, A.B. 1932. The Polyzoa, with a note on an associated hydroid. Scientific Reports of the Great Barrier Reef Expedition 1928-1929 4(12): 399-458
Haswell, W.A. 1881. On some Polyzoa from the Queensland coast. Proceedings of the Linnean Society of New South Wales 5: 33-44
Hayward, P.J. 1992. Some Antarctic and sub-Antarctic species of Celleporidae (Bryozoa, Cheilostomata). Journal of Zoology, London 226: 283-310
Hayward, P.J. & Thorpe, J.P. 1988. New genera of Antarctic cheilostome Bryozoa. Cahiers de Biologie Marine 29: 277-296
Hincks, T. 1881. Contributions towards a general history of the marine Polyzoa. VI. Polyzoa from Bass's Straits. Annals and Magazine of Natural History 5 8: 1-14, 122-128
Hincks, T. 1884. Contributions towards a general history of the marine Polyzoa. XIII. Polyzoa from Victoria and Western Australia. Annals and Magazine of Natural History 5 13: 363-369; 14: 276-285
Livingstone, A.A. 1928. Bryozoa from South Australia. Records of the South Australian Museum (Adelaide) 4(1): 111-124
Macgillivray, P.H. 1869. Descriptions of some new genera and species of Australian Polyzoa; to which is added a list of species found in Victoria. Transactions and Proceedings of the Royal Society of Victoria 9: 126-148
Macgillivray, P.H. 1879. Polyzoa. 21-40, pls 35-39 in McCoy, F. (ed.). Prodromus of the Zoology of Victoria Decade 4. Melbourne : George Robertson.
Macgillivray, P.H. 1880. Polyzoa. 27-52, pls 45-49 in McCoy, F. (ed.). Prodromus of the Zoology of Victoria Decade 5. Melbourne : George Robertson Vol. 5.
Macgillivray, P.H. 1883. Descriptions of new or little-known Polyzoa. Part 2. Transactions and Proceedings of the Royal Society of Victoria 19: 130-138
Macgillivray, P.H. 1895. A monograph of the Tertiary Polyzoa of Victoria. Transactions of the Royal Society of Victoria ns 4: 1-166
Maplestone, C.M. 1910. Observations on Parmularia obliqua and a fossil species. Transactions of the Royal Society of Victoria 23: 42-43
Rogick, M.D. 1955. Genus Emballotheca Levinsen 1909. Transactions of the American Microscopical Society 74: 103-112
Wass, R.E. 1984. Bryozoans: coloured mats of the sea. pp. 212-213 in Mead & Beckett Publishing (eds). Reader's Digest Book of the Great Barrier Reef. Sydney : Reader's Digest.
Wass, R.E. & Yoo, J.J. 1983. Cheilostome Bryozoa from the Southern Australian Continental Shelf. Australian Journal of Marine and Freshwater Research 34: 303-354
Waters, A.W. 1913. The marine fauna of British East Africa and Zanzibar, from collections made by Cyril Crossland M.A.,B.Sc.,F.Z.S., in the years 1901-1902. Bryozoa - Cheilostomata. Proceedings of the Zoological Society of London 1913: 458-537
History of changes
| Published | As part of group | Action Date | Action Type | Compiler(s) |
|---|---|---|---|---|
| 05-May-2015 | LANCEOPORIDAE Harmer, 1957 | 04-May-2015 | MODIFIED | |
| 25-Mar-2014 | BRYOZOA Ehrenberg, 1831 | 25-Mar-2014 | MODIFIED | Dr Robin Wilson (NMV) Elizabeth Greaves (NMV) |
| 12-Feb-2010 | (import) |