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Systematics , Anatomyandboringmecha PROC. R. SOC. VICT. vol. 96, no. 3, 113-125, Septem ber 1984 SYSTEMATICS, ANATOMY AND BORING MECHANISMS OF THE ROCK-BORING MYTILID BIVALVE BOTULA By B. R. W ilson and R ichard T a it M useum of Victoria, 285-321 Russell Streel, M elbour ne, Victoria, Australia 3000 A bstract : The mytilid Botu/a is shown to contain a single species, B. fusca Gmelin, which is widespread in tropical and sub-tropical waters of t he Indian, Pacific and western Atlantic O ceans. She ll m orphology and anatom y of this species are describe d. It is siphonate and bores in calcareous rocks an d dead corals by m eans of non-acid secretions from la rge boring glands in the m antle folds anteriorly an d posterodorsally. Its nearest living relatives are t he species of Lilhophaga s.s. and the genus is placed in the Lithophaginae along with Adula, Lilhophaga, Leiosolenus and Fungiacava. Recent classifications place the mytilid Botula M orch Natural History, Paris. NMV —Museum of Victoria, 1853 in the Modiolinae (e.g. Soot-Ryen 1969, Keen Melbourne. SAM —South Australian Museum, 1971). Other genera placed in that group are non- Adelaide. SDNHM —San Diego Natural History siphonate byssally attached ‘nestlers’ while Botula is a Museum, San Diego. WAM —Western Australian siphonate, rock-borer. Lamy (1937) and other author s M useum, Perth. place Botula in the Lithophaginae. This paper reports results of an anatomical study of Botula fusca, under­ taken to determ ine the affinity of the genus, to re view its SYSTEMATICS species taxonomy, and to determine the nature of th e Class B ivalvia boring mechanism. Family M ytilidae Genus Botula M orch 1853 METHODS 1853 Botula M orch, p. 55. 1939 Iredale, p. 414. (Type species: Living specimens of Botula fusca were collected and Botulopa Botulopa studied at the Heron Island M arine Research Station , in silicula infra Iredale, 1939 (= M. fusca herein)) the Great Barrier Reef Marine Park, Capricorn Zone. T ype species : Mytilus fuscus Gmelin 1791 by subse­ Supplementary preserved specimens were examined quent designation of Dali 1898. from several other Queensland and W estern Australia n D ia gnosis : Shell cylindrical, inflated, arcuate, sm ooth localities. For com parative purposes sections were made or finely concentrically striate, rather thin, white with a of the pallial boring glands of Leiosolenus lessepsianus, uniform thin, smooth, glistening brown periostracum . also collected at H eron Island. Umbos terminal, prosogyrate conspicuously incurved; Isotonic M gCh was used as an anaesthetic prior to hinge-line and ligament short, ligament opisthodeti c, dissection. Anatomical drawings were done from the parivincular; margins smooth except for fine vertic al anaesthetised specimens, free-hand or using a camer a- striae beneath and immediately posterior to the lucida. Specimens for histological work were fixed in ligament. Bouin’s solution, stored in 70% ethanol, routinely pro­ Incurrent and excurrent siphons long but usually cessed into paraffin blocks and sectioned at 5 /x. Sections divided; with a muscular septum within the tubular ex­ were taken in a plane perpendicular to the mantle e dge current siphon and a prom inent digitate ventral sip honal (transversely across the boring gland) and stained with branchial membrane. Ctenidia filibranchiate and M ayer’s haemotoxylin and eosin, M allory’s triple stain heterorhabdic, eleutherorhabdic. Boring glands pres ent or W iegert’s iron haemotoxylin. The periodic in the inner mantle folds anteriorly and postero­ acid —Schiff reaction (P.A .S.) in conjunction with Al- dorsally. Posterior siphonal retractor muscles and scars cian Blue (pH 2.35) was used as a test for acid or neutral present; anterior pedal retractors attach close to the mucopolysaccharides (M owry 1956). margin of the hinge just below the umbos; posterior Shells of the specimens on which the anatomical pedal-byssal retractors small. study is based are deposited in the M useum of Victo ria Mussels which bore in dead corals, coral-rock or Melbourne, the Australian Museum Sydney, and the other calcareous rocks. Western Australian Museum Perth. Abbreviations for respository institutions are as follows: AM —A ustra lian R emarks : M orch assigned to Botula only the species Museum, Sydney. ANSP—Academy of Natural arenaria M eusch. (with vagina Lamarck, and castaneus Sciences of Philadelphia, Philadelphia. BM(NH) — ‘Rum phius’ Gray (non Say) listed as synonyms) and British Museum (Natural History), London. fusca Gmelin (with M ytilus brunneus Solander, Modiola CAS —California Academy of Sciences, San Francisco. cinnamomea Lamarck and M. favanii Potiez and LACM —Los Angeles County Museum, Los Angeles. M ichaud listed as synonyms). Subsequently Dali (189 8) MCZ —Museum of Comparative Zoology, Harvard selected fusca Gmelin as type. In these authors’ opinion University, Boston. MNHN —National Museum of arenaria, a large, thin-shelled, siphonate mytilid which 113 114 B. R. WILSON AND RICHARD TAIT burrows in soft substrates of the central Indo-Pacific See Table 1 for details of type specimens. region, is not congeneric with fusca, although its correct D istr ibution : W idespread in the tropical and sub­ generic affinity is yet to be determ ined. tropical seas of the Indian, Pacific and western Atlantic Botula most resembles Lithophaga s.s. in that the Oceans. Apparently absent from the tropical eastern shell has a strong periostracum but no secondary A tlantic. Figure 7 is based on verified m useum spec imen calcareous accretions, there are prom inent plicate mem­ records and selected literature records. branes in the roof of the mantle cavity, and boring is aided by chemical secretions from anterior and dors al H abitat : Botula fusca is a boring mytilid inhabiting boring glands. It is easily distinguished from that genus dead corals, coral rocks and other calcareous rocks . It is by the short, arcuate shell form, the large, strong ly common in the intertidal zone and has been collecte d incurved umbos, and the divided posterior siphons. from suitable substrates to depths as great as 291-190 fathom s (off Hawaii, see Dali, Bartsch & Rehder 193 8). Botula fusca (Gmelin 1791) It is often extraordinarily abundant in coral rocks and Figs 1 A -P, 2 B-F, 3-7 the coral-rock platform of the reef-crest zone, sometimes most commonly with the burrows opening on 1785 Mytilus cinnamominus Chem nitz, pi. 82, fig. 731. the underside of the rocks. The species may also be (Refers to Lister, 1687, fig. 197.) (Type locality: abundant burrowing in sloping beach-rock relatively Jam aica.) (Non-binomial publication.) high in the intertidal zone, e.g. along the souther n shore 1791 M ytilus fuscus Gmelin, p. 3359. (Refers to Lister, of Heron Island (Queensland) below the research sta ­ 1687, fig. 197.) (Type locality: Jam aica.) tion. It is also known to burrow in the shells of o ther 1793 Mytilus cinnamomeus Schreibers, p. 293. (Refers molluscs such as Strombus gigas (Chemnitz 1785) and to Chem nitz, 1785, pi. 82, fig. 731.) oysters (Berry 1959). In northern A ustralia it is c ommon 1807 Modiolus cinnamomeus Link, p. 147. (Refers to boring in massed shells of rock-oysters (Saccostrea) high Chem nitz, 1785, pi. 82, fig. 731.) in the intertidal zone. 1819 Modiola cinnamomea Lamarck, p. 114. (Type Gohar and Soliman (1963) have described and locality: l’lsle de France (Mauritius, Indian figured burrows of this species (identified as Modiolus Ocean), leg. M . Desetangs.) cinnamomeus Bruguiere) from the Red Sea. The burrow 1819 M odiola silicula Lam arck, p. 114. (Type locality: lacks a calcareous lining and is differentiated into two ‘Nouvelle Hollande’ probably collected by Peron parts. The outer or siphonal part is roughly dumb-b ell during the Baudin Expedition at Shark Bay, shaped in cross-section and it widens distally. The inner W estern Australia.) part, which is occupied by the shell, is kidney-sha ped 1844 Modiole favanni Potiez & M ichaud, p. 130, pi. 54, like the shell. fig. 9. (Type locality: ‘Exotique’.) 1892 Lithodomus projectans Tate, p. 130, pi. 1, fig. 1. D escr iption : Shell. W hite to pale brown, covered by a (Type locality: ‘Port Darwin’.) uniform tan to dark-brown or almost black glossy 1938 Botula hawaiensis Dali, Bartsch & Rehder, 1938, sm ooth periostracum . Cylindrical-arcuate, anterior and p. 59, pi. 12, figs. 1-4. (Type locality: ‘dredged by posterior ends broadly rounded; prosogyrate umbos the U.S. Bureau of Fisheries Steamer “A lbatross” large and term inal, sometimes conspicuously projecting at Station 3845 off the south coast of M olokai and incurved. Ligament opisthodetic, parivincular, (Hawaii) in 60-64-0 fathom s on sand, pebble and short and thick, resilium entire, sub-ligamental ridge shell bottom ; bottom tem perature 71 °F.’) lacking; hinge edentulous, margins smooth except fo r 1938 Botula laysana Dali, Bartsch & Rehder, 1938, p. very fine, oblique striae on the dorsal margin bene ath 60, pi. 12, fig. 5. (Type locality: ‘dredged by the and immediately behind the ligament. Exterior smoot h U.S. Bureau of Fisheries Steamer “Albatross” at except for weak concentric growth striae and, in so me Station 3936, near Laysan Island (Hawaiian cases, series of concentric ridges presumably repre ­ Islands) in 79-130-0 fathom s on small broken shell senting successive growth phases (Fig. ID , G; cf. and coralline bottom ; bottom tem perature 68°F.’) Lithophaga teres and L. antillarum, W ilson 1979, p. 1939 Botulopa silicula infra Iredale, p. 415, pi. 6, fig. 440, 448). Scars of anterior and posterior retracto rs 26. (Type locality: Low Isles, Queensland.) weak but usually discernible, scar of siphonal retr actor 1959 Botula cylista Berry, p. 108. (Type locality: Punta very weak but discernible. Cameron, M azatlan, Sinaloa, Mexico.) A natom y (see also Pelseneer 1911, Soot-Ryen 1955, Fig.
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