Family SEPIIDAE

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Family SEPIIDAE

Compiler and date details

July 2001 - C.C. Lu, National Chung Hsing University, Taichung, Taiwan

Introduction

Members of the family Sepiidae Keferstein, 1866 are common on the continental shelf and upper slope of the temperate to tropical waters of the Old World. They are found from Japan, China, the Philippines and Indo-Malaya to Australia, eastward to Fiji and westward around the Indian Ocean to the Eastern Atlantic as far north as the North Sea. The family does not occur around the Americas. In the Pacific the easternmost records are from the Marshall Islands and Fiji; no sepiids are known to occur in New Zealand waters.

The family comprises some 100 nominal species in three genera: Sepia, Metasepia and Sepiella (Lu & Reid 1997; Khromov et al. 1998; Reid & Lu 1998, Reid 2000). Thirty-three species representing all three genera have been reported from Australian waters (Lu & Reid 1997; Lu 1998; Reid 1998; Reid & Lu 1998; Reid 2000).

The study and naming of cuttlefish dates back to Linnaeus who named the common cuttlefish of Europe Sepia officinalis in 1758. Subsequently, many authors have contributed to the description and naming of members of this family, notably, Lesueur (1821), Férussac and d'Orbigny (1835–1848), Gray (1849), and more recently Voss (1963), Adam (in Adam & Rees 1964), Roeleveld (1972), Roeleveld and Liltved (1985), and Khromov (1987a, 1987b, 1987c; 1990). Despite the numerous published papers dealing with the cuttlefish taxonomy, detailed taxonomic revisions are few. Rochebrune (1884) attempted to monograph the family but the classification he proposed was largely refuted by Adam and Rees (1964). Roeleveld's (1972) monograph clarified the Sepiidae of South African waters. Khromov (1987c, 1990) reported his work on the classification and phylogeny of Sepiidae. Recently, Khromov et al. (1998) and Lu (1998) have provided synoptic reviews of the family.

The first descriptions of cuttlefish from Australian waters were published in the first half of the 19th century (Lesueur 1821; Gray 1849), but the taxonomy and biology of the Australian cuttlefish are as yet poorly known. Brazier (1892) listed 14 species of Sepia from Australia, among them several names from the South African fauna. Berry (1918) described three species of Sepia from southern Australia. Between 1926 and 1954, Iredale (1926a, 1926b, 1940, 1954) and Cotton (1929, 1931, 1932) described and named 29 and seven cuttlefish taxa, respectively. All were based solely on the sepions; the soft parts of the animals were not described. Adam (1979) reported on the cuttlefish collection in the Western Australian Museum; he described 18 species of Sepia and clarified the status of many of Iredale and Cotton's taxa. Lu & Phillips (1985) and Lu (1998), incorporating Adam's (1979) findings with new data, summarised the status and distribution of the Sepiidae from Australian waters. Roper & Hochberg (1988) after studying the colour pattern and locomotion of Sepia (Metasepia) pfefferi at Lizard Island elevated the subgenus Metasepia to generic status. Lu (1998) recognised 26 species in all three genera, Sepia, Metasepia and Sepiella, from Australian waters; three known only from cuttlebones were designated as of uncertain status. Six further species have been added to the list since the preparation of that paper (Lu & Reid 1997; Reid & Lu 1998; Reid 2000).

Recent work reveals that of the 33 species (excluding the two species known only from sepions), 26 are considered to be endemic to Australian waters (Lu & Reid 1997; Lu 1998; Reid 1998; Reid & Lu 1998). The figure strongly supports Adam & Rees (1966) assertion that worldwide, the Australian sepiid fauna is among the richest in endemic species.

The normal life-span of the common European cuttlefish Sepia officinalis varies from 18 to 24 months although some individual male may live longer. Mass mortality at the end of the spawning period occurs on the Atlantic Coast; no comparable intensity has been reported in the Mediterranean. In Australia, mass mortality of Sepia apama has been observed in July/August off the coast of New South Wales (Battam, pers. comm.). Examination of two of the individuals involved revealed that the male (382 mm mantle length (ML), 6.2 kg) was mature and the female (300 mm ML, 2.4 kg) immature; the stomachs of both individuals were empty. The cause of such mortality is uncertain.

In Sepia officinalis, a positive correlation has been found between temperature and early growth rate, and between maximum size and life span. Negative correlations are reported between maximum size or age and early growth rate, and between temperature and life span (Richard, cited in Boletzky 1983). Pascual (1978), on the basis of rearing experiments, concluded that faster growth at higher temperatures is due to higher food intake; that the efficiency of food conversion is independent of temperature. The growth rate of Sepia apama has been estimated to be 10 mm per month, the animals taking 10 to 12 months to reach adult size (Bell 1979). A large specimen, with ML 460 mm, was estimated to be about four years old.

Cuttlefish may attain sexual maturity at very different sizes. Mangold-Wirz (1963) reported that male Sepia officinalis can be fully mature as small as 6–8 cm ML, while individuals as large as 10 cm ML may still be immature; females also exhibit the same phenomenon, maturing at sizes varying from 11 to 25 cm ML. The same phenomenon has been observed in Sepia pharaonis from the Gulf of Carpentaria where the mature males range from 4.7 cm ML to 14.5 cm ML and mature females range from 14.1 cm ML to 17.4 cm ML; the largest immature female measured 14.3 cm ML (Dunning et al. 1994; Lu, unpublished). The factors responsible for this variation in body size at maturity were found to be the combined effect of temperature and light (Richard, cited in Boletzky 1983).

In Sepia officinalis spawning occurs soon after mating. Eggs are laid in regular intervals of 2–3 minutes over several hours. The egg is 2.5–3 cm in length, 1.2–1.4 cm in the greatest diameter. Each is fixed to any oblong object by means of a ring-shaped basal structure. The ring is produced by the animal which uses its arm tips to draw the gelatinous envelope of the egg into a pair of processes and then winds them round the supporting object so that they stick together (Boletzky 1983). A female may empty her ovary of mature eggs within a few days. Mangold-Wirz (1963) also found that the total number of eggs in an individual varies from 500 to over 1000, depending on the size of the animal. The duration of embryonic development varies with temperature and ranges from 40–45 days at 20°C to 80–90 days at 15°C. Hatchlings of Sepia officinalis have ML 6–9 mm and are similar to the adult in overall features.

Predators on cuttlefishes are many and include the short-finned pilot whale, Globicephala macrorhyncha in South Africa (Roeleveld in Clarke 1986), the pygmy sperm whale, Kogia breviceps and the dwarf sperm whale Kogia simus in South Africa (Ross 1979 cited in Clarke 1986), and Risso's dolphin, Grampus griseus in the English Channel (Clarke & Pascoe 1985). In Australia, they are known to be preyed upon by the long-finned pilot whale Globicephala melas and the Australian fur seal Arctocephalus pusillus doriferus in Tasmania (Gales et al. 1992, 1994).

The prey of Sepia species consists of a wide variety of organisms. Najai & Ktari (1974 cited in Boletzky 1983) found decapods including Penaeus species, isopods such as species of Sphaeroma, Cymodocea, copepods and ostracods, bivalves, gastropods and pteropods, octopods and decapod cephalopods, polychaetes and nemerteans and fishes in the stomach contents of Sepia officinalis from the coast of Tunisia. Little is known about food of cuttlefish in Australian waters, however, crustacean remains have been found in the stomachs of large numbers of individuals of Sepia pharaonis, S. elliptica and S. smithi from the Gulf of Carpentaria (Lu 2001).

Excluded Taxa

Misidentifications: SEPIIDAE: Sepia aculeata Van Hasselt in Férussac & d'Orbigny, 1848 [occurrence in Queensland is unconfirmed; Lu, personal communication, regards this species excluded from the Australian fauna; Sepia aculeata occurs in the Indo-Pacific south to Java] — Brazier, J. 1892. Catalogue of the Marine Shells of Australia and Tasmania. Pt I. Cephalopoda; Pt II. Pteropoda. Sydney : Australian Museum Catalogue Vol. 15 42 pp. (listed species from Australian waters); Lu, C.C. & Phillips, J.U. 1985. An annotated checklist of Cephalopoda from Australian waters. Occasional Papers of the Museum of Victoria 2: 21-36 (removed species from Australian distribution); Lu, C.C. 2001. Cephalopoda. pp. 129-308 in Wells, A. & Houston, W.W.K. (eds). Zoological Catalogue of Australia. Vol. 17.2 Mollusca: Aplacophora, Polyplacophora, Scaphopoda, Cephalopoda. Melbourne : CSIRO Publishing, Australia xii 353 pp. [Date published 3 July 2001] [185] (listed the species from Australian waters)

Diagnosis

The cuttlefish of this speciose family are small to medium sized (although several species reach 50 cm in mantle length), and bear a characteristic calcareous sepion (cuttlebone). The sepion is porous, finely laminated and is located on the dorsal side of the mantle underneath the skin. The spine at the posterior end of the sepion may be absent in some species. The mantle is broad, or slender, robust, elongate oval to circular in outline and slightly flattened dorso-ventrally. The head is robust with prominent eyes that are covered by a transparent membrane; the eyelid has a conspicuous secondary fold. The funnel-mantle locking apparatus is curved to angular in shape. The fins are narrow, lateral to the mantle and occupy almost whole length of the mantle. The posterior end of the fins is not connected to the mantle, but forms a free lobe. The eight arms are equipped with two to four rows of suckers. The tentacles can be retracted into a pocket on the ventro-lateral sides of the head. The tentacular clubs bear four to eight or more (may be more than 20) rows of suckers, depending on the species.

The calcareous shell (sepion) lies under the skin on the dorsal side of the mantle, running almost the entire length of the mantle. The shape of the sepion ranges from lanceolate, oval to rhomboidal. The dorsal side of the sepion is a calcareous plate called dorsal shield. Ventrally the sepion is a series of numerous thin, narrow, oblique septa which are supported by numerous transverse calcareous rods. A spine is present at the posterior end of the sepion in most species, with the exception of Sepiella species and a few species of Sepia.