Class CEPHALOPODA Cuvier, 1795

Cephalopoda

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Class CEPHALOPODA Cuvier, 1795

Cuttlefish, Nautilus, Octopus, Squid

Compiler and date details

2024 - updated (part) A.L. Reid, CSIRO, Australia.

2007 - Sepiida updated by A.L. Reid, University of Wollongong, Australia

2003, 2007 - updated (part), Mark Norman, Museum Victoria, Melbourne

30 June 2000 - C.C. Lu, National Chung Hsing University, Taichung, Taiwan

Introduction

The Class Cephalopoda Cuvier, 1798 comprises three subclasses, Ammonoidea, Nautiloidea, and Coleoidea. All ammonoids are extinct. The Nautiloidea comprises 15 orders; only the order Nautilida is still extant, and within it only the family Nautilidae, and two genera, Nautilus and Allonautilus. All other living cephalopods belong to the Coleoidea. Extant coleoids are divided into six orders and 47 families (WoRMS, 2024). Forty-four families, represented by approximately 240 species, have been found in Australian waters.

Life histories are unknown for most cephalopods in the Australian fauna and those we know at all are fragmentary at best (Boyle 1983, 1987) and based largely on work done on relevant groups outside Australia.

Coleoid cephalopods are often described as having a live-fast-die-young lifestyle, with most species living for about a year, and a repeated spawning in a short period at the end of the individual’s, life. This knowledge of life history strategies is, however, biased toward commercially exploited epipelagic and inshore species, especially from the family of flying squids (Ommastrephidae). While this strategy may apply to many inshore species, scientific evidence is accumulating to suggest that members of most oceanic squid species do not, in fact, have a short lifespan of one year or less and repeated spawning at the end of the individual’s, life (Hoving 2012). The same is true for species that inhabit colder and, or, deep waters with consequently slower growth rates (Collins and Rodhouse 2006, Hoving et al. 2014). However, life spans generally range from a few months for tiny species to fewer than five years at most (Hanlon et al. 2018). Nautilids in aquaria are known to live 15 years or more.

Most species are semelparous, although exceptions do occur. Cephalopods are dioecious, and many species show sexual dimorphism in size or body proportions, extremes being some pelagic incirrate octopods such as Argonauta and Ocythoe in which adult males are dwarfs. Cephalopod development is direct: hatchlings of species with large eggs resemble miniature adults, and hatchlings of species with small eggs undergo only gradual change in body proportions (Young & Harman 1988; Sweeney et al. 1992). Most cephalopods are active, opportunistic carnivores. Nautilidae species, however, feed close to the bottom, and may scavenge. Coleoids prey upon crustaceans, fishes and molluscs including cephalopods. Many examples of cannibalism are known. Other prey include echinoderms and polychaetes (Nixon 1987).

Rapid growth in cephalopod fisheries has occurred worldwide in recent years. Cephalopods form only a minor component of the Australian fisheries industry at present, with most consumption consisting of squid as calamari, but recently the demand for octopus has increased leading to fully functioning fishers in Tasmania and Western Australia with developing fisheries in Victoria and South Australia. However, this is likely to change if the fin fisheries in Australia and the rest of the world continue their present collapse and continue to expand into lower trophic level species as new protein sources for vast human populations are sought. Pressure on Australian stocks will undoubtedly increase. Their management is compromised not only because of remaining gaps in our understanding of species boundaries, but also in our knowledge of cephalopod life histories and ecosystem functions. In order to manage sustainable cephalopod fisheries, there needs to be a greater understanding of population dynamics and the role of cephalopods in ecosystems. This needs to become a high research priority as relatively few such detailed studies have been undertaken on Australian species.

Population recognition and size estimates are also essential elements in the analysis of the conservation status of species under the International Union for the Conservation of Nature (IUCN) Red List Criteria (www.iucnredlist.org) and under the International Trade of Endangered Species (CITES) listing criteria for protecting species that are in international trade (www.cities.org). There is little data available on population sizes and density of Australian cephalopods to enable a preliminary baseline for such assessments.

Genetic studies of population structuring and dispersal patterns are also important areas for future research in addition to more extensive use of molecular identification tools to underpin any expansion of Australian cephalopod fisheries to be applied to questions relating to correct labelling, authenticity, traceability and import and export control.

In addition to their direct contribution to fisheries production, coleoid cephalopods play an important role in the ecology of the ocean. They are important prey for marine mammals such as toothed whales, many species of seals and seabirds, and large fishes such as tunas and sharks. In turn they consume large quantities of crustaceans, cephalopods and fishes. Their indirect importance to marine fisheries is considerable.

Traditionally, extant Coleoidea have been divided into three orders: Sepioidea, Teuthoidea, Vampyromorpha and Octopoda (G.L. Voss 1977, 1988a, 1988b). More recently, the following Orders have been recognised: Bathyteuthida, Idiosepida, Myopsida, Oegopsida, Sepiida, Sepiolida, and Spirulida in the Superorder Decapodiformes and Octopoda and Vampyromorpha in the Superorder Octopodiformes (WoRMS, 2024). All are represented in Australian waters.

Taxonomic description of coleoids from Australian waters dates back to Lesueur (1821), however, due to the lack of detailed descriptions and illustrations, these early species remain unrecognisable. Quoy & Gaimard (1832) reported the zoological results of Dumont d'Urville's voyage aboard l'Astrolabe (1826–1829) and described four species from south-eastern Australia; Gray (1849) and Hoyle (1886) both described many species from Australia. In 1892, Brazier published the first comprehensive list of Australian cephalopods, including 20 octopod species, 19 cuttlefish species, eight myopsid squid species, five oegopsid squid species and four Nautilus species.

Twentieth Century works include Berry's (1918) report on the cephalopod collections made by the F.I.S. Endeavour in south-eastern Australia between 1909 and 1914, with 13 species, eight of them new. The next major reports were those of Iredale (1926, 1940, 1954), and Cotton's (1929, 1931, 1932) studies of Australian cuttlefish, among them descriptions of 35 new species or subspecies. In 1945, Allen published the results of her studies of the planktonic cephalopod larvae from eastern Australian waters; she reported 21 species, including two new species and several new records for Australia. Adam's (1979) report on the Sepiidae in the collections of the Western Australian Museum was the next significant study. He reported the species of Sepia from Western Australia, among them three new species; he also included two species from eastern Australia.

Interest in cephalopod fisheries, and especially in squid fisheries, in Australia in the late 1970's stimulated the increased research effort on Australian cephalopods. A consequence of this increased interest and awareness of the importance of cephalopods in the marine ecosystem, resulted in active expansion of cephalopod collections in institutions. Since 1979, many papers on Australian cephalopods have been published. In 1985, Lu and Phillips published an annotated checklist of cephalopods from Australian waters, the first comprehensive checklist since Brazier's 1892 checklist. These authors listed 222 nominal species, in 35 families and 62 genera, and including four Nautilus species, 80 sepioid species, 15 myopsid squid species, 78 oegopsid squid species, 44 octopod species and the monotypic Vampyroteuthis infernalis. Many more articles on the Australian coleoid fauna have since been published.

BIOGEOGRAPHY
The Australian marine fauna is considered to comprise two distinct latitudinal elements separated by zones of intermixing. These faunal regions were designated by Wilson & Gillet (1971) as the Northern Australian Region, Southern Australian Region and Eastern and Western Overlapping zones. The Northern Australian Region is part of the Indo-west Pacific Faunal Region and is characterised by high species diversity and large numbers of species with wide distributions in the Indian and Western Pacific Oceans. The fauna of the Southern Australian Region is characterised by low diversity and high species endemicity. The Overlap zones are long stretches of coast between the Northern and Southern Regions; there is a gradual replacement of tropical forms with temperate forms, and some local endemism (Wilson & Allen 1987).The Australian neritic coleoid fauna generally conforms to this distributional pattern.

The distributional pattern of pelagic cephalopods is less clear-cut. Although data are only preliminary, a latitudinal gradient in species diversity, such as observed in the Eastern North Atlantic Ocean (Clarke & Lu 1974, 1975; Lu & Clarke 1975a, 1975b), is not evident. This impression is supported also by Nesis' (1979) interpretations of pelagic cephalopod data, upon which basis he classified the pelagic realm of the Australian-New Zealand region into equatorial, southern subtropical, peripheral, notal and Antarctic zones. Only the first four zones are present in the Australian Fishing Zone (AFZ)— the Antarctic zone lies outside the AFZ. The equatorial and the southern subtropical zones encompass the entire eastern, western and northern waters, while the peripheral zone encompasses the Great Australian Bight region, including Tasmania. The notal zone comes close to the south-western corner of Australia but largely lies outside of AFZ (Nesis 1979: fig. 3).

The broad distributional ranges of many species have resulted in a diffuse pattern with respect to the latitudinal gradient in species diversity. The southward flow of both the warm Leeuwin Current in the west and the warm East Australian Current in the east (see Bunt 1987) also are contributing factors. The Leeuwin Current flows eastward around Cape Leeuwin, bringing warm, tropical water to the Great Australian Bight and supporting a tropical marine fauna in the Bight (Maxwell & Cresswell 1981). The East Australian Current flows east at around 33°S, where warm-core anticyclonic eddies are formed by pinching off the East Australian Current and continue southward to as far south as 40°S. Brandt (1983) suggested that these eddies may be responsible for the large-scale patchiness in pelagic distribution in the western Tasman Sea. The eddies may carry tropical species beyond their normal range. A juvenile Spirula spirula taken in the zooplankton off south-eastern Tasmania (44°05.6'S 147°58.6'E) and juvenile Sthenoteuthis off Bass Strait (Dunning 1988c) may be examples (Lu, unpublished data).
The effects of climate change, resulting in the more southward movement of the East Australian Current is having an effect on species distributions, with paralarval cephalopods able to survive in regions beyond their former range due to increasing water temperatures. The discovery of Octopus tetricus in Tasmania is an example. This species was previously restricted to mainland Australia.

FOSSIL HISTORY
Records of Australian coleoid fossils are few. The family Bolemnidae is represented by several Dimitobelus species from the Cretaceous in South Australia (Ludbrook 1966). Fossils of Spirulirostra curta (Family Spirulirostridae) have been collected from the Muddy Creek Formation (Middle Miocene, Balcombian) in Torquay (NMV collection), and of Notosepia cliftonensis (Family Sepiidae) from the Balcombian in Dingley and Hamilton, Victoria (NMV collection). Muensterella tonii Wade of the extinct order of Kelaenida Starobogatov and fossil vampyromorphs, Boreopeltis soniae Wade and Trachyteuthis willisi Wade of the families Plesioteuthidae and Trachyteuthidae, respectively, have been reported from Queensland (Wade 1993).

Acknowledgements

The project was initially supported by a grant to C.C. Lu from the Australian Biological Resources Study (ABRS) which is gratefully acknowledged. C.C. Lu thanks Alice Wells and Keith Houston for editorial advice for early drafts and good humour and patience during the long drawn out period of preparation and is also indebted to Malcolm Dunning who patiently read the manuscript, noted many errors and provided many suggestions.

Lu also acknowledges Curators and Collection Managers of cephalopod collections in all Australian museums, as well as Mike Sweeney of the Division of Mollusks of the Smithsonian Institution, tirelessly and patiently responded to all my requests for loans of specimens or information.

He thanks Val Hogan, Frank Job and Sandra Winchester of the Library of the Museum of Victoria who tirelessly assisted with location of many obscure references needed for the project and obtained necessary references through numerous interlibrary loans. Without their help it would have been impossible to complete the project.

Lu is indebted to my many friends and colleagues, notably, Sue Boyd, Malcolm Dunning, Robyn Ickeringill, Mandy Reid, and Chris Rowley, who have answered many requests for information on specimens or literature.

Introductory remarks for each family in this section are largely abbreviated versions of the family treatments in Chapter 13, Subclass Coleoidea, of the Fauna of Australia Volume 5. Lu thanks his good friend and colleague, Malcolm Dunning for agreeing to let him rely so heavily on the teuthid section that he authored.

In the early stages of the project, Margaret Blackburn and later Timothy Stranks were employed to enter data. Their efforts are acknowledged.

Recent edits have been made by Dr Mandy Reid based on name updates, taxonomic revisions and heavily relied on her publication ‘Cephalopods of Australia and sub-Antarctic Territories’ CSIRO Publishing: Melbourne. Julian Finn and Mark Norman have also contributed to recent updates. Some of the information, particularly regarding biology, behaviour and fisheries is now a little outdated, so researchers are referred, where possible, to more recent literature for updates.

Database Notes

The information on the Australian Faunal Directory site for the Cephalopoda is derived from the Zoological Catalogue of Australia database compiled on the Platypus software program. It incorporates a few minor changes made to the work published on 2 July 2001 as

Taxa are arranged alphabetically. Synonyms are arranged in chronological order and only those relevant to Australian records are considered.

All type specimens housed in Australian institutions have been examined for verification of data. Information on types deposited in museums outside Australia is based mostly on published data from the original publications or derived from the following type catalogues: Kristensen & Knudson (1983), Lu et al. (1995), and Sweeney & Roper (1998). Several recent publications that have been very useful and have been drawn upon heavily are Lu (1998), Lu & Dunning (1998) and Sweeney & Roper (1998).

No common names are given in this work, but common names of cephalopods are used in the Internet database, CephBase.

Minor changes were made in January 2007, following the publication, Groves et al. (2006).

Limital Area

Distribution data in the Directory is by political and geographic region descriptors and serves as a guide to the distribution of a taxon. For details of a taxon's distribution, the reader should consult the cited references (if any) at genus and species levels.

Australia is defined as including Lord Howe Is., Norfolk Is., Cocos (Keeling) Ils, Christmas Is., Ashmore and Cartier Ils, Macquarie Is., Australian Antarctic Territory, Heard and McDonald Ils, and the waters associated with these land areas of Australian political responsibility. Political areas include the adjacent waters.

Terrestrial geographical terms are based on the drainage systems of continental Australia, while marine terms are self explanatory except as follows: the boundary between the coastal and oceanic zones is the 200 m contour; the Arafura Sea extends from Cape York to 124 DEG E; and the boundary between the Tasman and Coral Seas is considered to be the latitude of Fraser Island, also regarded as the southern terminus of the Great Barrier Reef.

Distribution records, if any, outside of these areas are listed as extralimital. The distribution descriptors for each species are collated to genus level. Users are advised that extralimital distribution for some taxa may not be complete.