Overview
Brief Summary
Introduction
The Australian Giant Cuttlefish (Sepia apama) is the largest cuttlefish species in the world, with a maximum recorded size of 520 mm mantle length (ML) and 6.2 kg weight . It is endemic to Australian waters, with a distribution reported to extend across temperate southern Australia from southern Queensland to Point Cloates in Western Australia, and including northern Tasmania. Its life span is 1 year, perhaps 2 in some cases. It occurs on rocky reefs, seagrass beds and areas of mud and sand to depths of 100m. This species is famous for its unique and very large spawning aggregation that occurs every austral fall (May - July) in northern Spencer Gulf, northwest of Adelaide.
References
Hall, K. C. and R. T. Hanlon. 2002. Principal features of the mating system of a large spawning aggregation of the giant Australian cuttlefish Sepia apama (Mollusca : Cephalopoda). Mar Biol 140: 533-545
Norman, M. R. 2000. Cephalopods, a world guide : Pacific Ocean, Indian Ocean, Red Sea, Atlantic Ocean, Caribbean, Arctic, Antarctic, ConchBooks, Hackenheim, Germany 318 pp.
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Interesting Facts
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Succinct
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Comprehensive Description
Description of Sepia apama
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Coloration
Sepia apama skin posesses a dense layer of pigmented chromatophore organs of 3 color classes: yellow, red, and black/brown, as well as underlying structural reflectors of 2 types: iridophores (creating iridescence in the skin ) and leucophores, which produce whiteness in their body patterns. They also have controllable skin papillae that can dramatically alter their appearance. The overall appearance of the animal - termed the body pattern - can change in less than a second due to direct neural control of the skin patterns. This is unique among all animals but is characteristic of cephalopods.
Reference
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Distribution
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UNESCO-IOC Register of Marine Organisms
http://www.marinespecies.org/aphia.php?p=sourcedetails&id=1318
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Jereb, P.; Roper, C.F.E. (Eds)(2005). An annotated an illustrated catalogue of cephalopod species known to date. Volume 1: Chambered nautilusses and sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes 4(1). FAO, Rome. 262p., 9 colour plates.
http://www.marinespecies.org/mollusca/aphia.php?p=sourcedetails&id=124589
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Sepia apama occurs in the southern half of Austrlia from Brisbane in the east to Sharks Bay in the west.
Reference
Norman, M. R. 2000. Cephalopods, a world guide : Pacific Ocean, Indian Ocean, Red Sea, Atlantic Ocean, Caribbean, Arctic, Antarctic, ConchBooks, Hackenheim, Germany 318 pp.
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Physical Description
Size
Physical Description
The giant cuttelfish is the largest cuttelfish species in the world. Sepia apama have eight sucker-lined arms and two retractable tentacles, which they use for capturing prey. Cuttlefish have a highly developed central nervous system and well developed complex eyes. They have a thick, internal calcified shell (the "cuttlebone") beneath an elongated muscular mantle. The mantle is expanded and contracted to expel water from the mantle cavity through the funnel. The mouth consists of a parrot-like beak, jaws, and a rasping tongue (a typical molluscan radula).
References
BBC Giant Cuttlefish Fact File
Hanlon, R.T. & J.B. Messenger. 1996. Cephalopod Behaviour. Cambridge University Press, 232pp.
Nixon, M. & J. Z. Young. 2003. The Brains and Lives of Cephalopods. Oxford University Press, 392pp
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Diagnostic Description
Formal Description
Diagnostic features: Adults very large. Sucker-bearing surface of tentacular club raised off of the stalk, attached only by a thin membrane; 5 suckers in rows across the manus, median suckers enlarged; swimming keel of the club extending along stalk a distance equal to the club length. Web between arms deep: equal to half of arm length between dorsal arms, two-thirds of arm length between lateral arms, absent between ventral arms. Three flat, semicircular, flap-like papillae posterior to each eye. Fins broad. Colour: deep maroon.
References
Roper, C. F. E., M. J. Sweeney and C. E. Nauen. 1984 FAO Species Catalogue Vol 3. Cephalopods of the World: An Annotated and Illustrated Catalogue of Species of Interest to Fisheries. FAO Fisheries Synopsis 125(3).
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Ecology
Habitat
Water temperature and chemistry ranges based on 5 samples.
Environmental ranges
Depth range (m): 1.5 - 71.5
Temperature range (°C): 14.137 - 17.433
Nitrate (umol/L): 0.094 - 2.747
Salinity (PPS): 35.417 - 35.773
Oxygen (ml/l): 5.358 - 5.626
Phosphate (umol/l): 0.139 - 0.312
Silicate (umol/l): 0.831 - 2.015
Graphical representation
Depth range (m): 1.5 - 71.5
Temperature range (°C): 14.137 - 17.433
Nitrate (umol/L): 0.094 - 2.747
Salinity (PPS): 35.417 - 35.773
Oxygen (ml/l): 5.358 - 5.626
Phosphate (umol/l): 0.139 - 0.312
Silicate (umol/l): 0.831 - 2.015
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.
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Population Biology
Population Structure
In general, the sex ratio in the populations is thought to be 1:1. However, there is a highly skewed male-biased operational sex ratio in the spawning aggregation (up to 11:1). This is the highest known among cephalopods. The change in the S. apama sex ratio during the spawning season (from more to less skewed towards males) suggests that males arrive before females at the start of the season. Therefore, males may not aggregate in response to a concentration of females, but rather at a particular location or habitat type. Recent genetic evidence (using allozyme electrophoresis, microsatellites and mitochondrial nucleotide sequences) has revealed three major genetically distinct groups of Sepia apama. It has been recommended that each of these groups be managed separately to preserve genetic diversity.
Reference
Hall, K. C. and R. T. Hanlon. 2002. Principal features of the mating system of a large spawning aggregation of the giant Australian cuttlefish Sepia apama (Mollusca : Cephalopoda). Mar Biol 140: 533-545
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General Ecology
Ecological Determinants/Niche
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Ecosystem Role
Sepia apama eat crustaceans and fish and have many predatorrs including: fur seals, sea lions, albatross, dolphins and sharks.
References
Battam, H. and W. A. Buttemer. 2000. Some aspects of energy assimilation and use in four southern albatrosses. Marine Ornithology 28.
Gales, R., D. Pemberton, C. C. Lu, and M. R. Clarke. 1993. Cephalopod diet of the Australian fur seal - variation due to location, season and sample type. Australian Journal of Marine and Freshwater Research 44: 657-671.
McIntosh, R. R., B. Page, and S. D. Goldsworthy. 2006. Dietary analysis of regurgitates and stomach samples from free-living Australian sea lions. Wildl Res 33: 661-669.
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Life History and Behavior
Behavior
Giant cuttlefish have the remarkable ability to dynamically change color and pattern for a wide variety of key behaviors involving camouflage and communication. Sepia apama is mainly active during the day and uses camouflage to hide among rock reefs and seaweed. Cuttlefish usually sit on or hover slightly above the bottom. Sepia apama can move slowly with stealth, swim or employ jet propulsion in bursts of surprising speed. This species preys on live fish, crabs and other crustaceans. All members of the Sepioidea use jets of ink to confuse foes during escapes. They are generally solitary animals (except when breeding) and curious about divers. Sepia apama skin posesses a dense layer of pigmented chromatophores, mainly yellow, red, black and brown, as well as a wide variety of iridophores (creating iridescence in the skin for camouflage and communication) and leucophores that produce whiteness in their body patterns. They also have controllable skin papillae that can dramatically alter their appearance.
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Night Camouflage
In addition to camouflaging themselves during daylight hours, Sepia apama uses adaptable night camouflage to conceal itself from nightime predators; animals in spawning aggregations have been shown to cease sexual signaling behaviors and become sessile and camouflaged during the night. Remarkably, the camouflaged body patterns at night are tailored to each microhabitat that the animal might rest in, providing evidence that their night vision is excellent and that night predators also have keen vision.
References
Hanlon, R. T., M. J. Naud, J. W. Forsythe, K. Hall, A. C. Watson, and J. McKechnie. 2007. Adaptable night camouflage by cuttlefish. Am Nat 169: 543-551
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General Camouflage
Cuttlefish are capable of showing dozens of body patterns for camouflage and these body patterns can be grouped into three categories: uniform/stipple, mottle, and disruptive. Aside from skin patterns, cuttlefish can augment camouflage with postural and 3-D skin texture: by contorting their arms in different postures that generally imitate nearby algae, and by sprouting spiky skin projections, called papillae, imitating the physical texture of the surrounding seaweed, rock or coral. The chromatophores and skin papillae can change about one second faster than any other animal. Both skin texture and color changes are directly controlled by the animal's brain. Messages enter the brain through the eye via the optic nerve, are processed in the CNS, and then the skin is changed by direct neural control of the pigmented chromatophores.
References
Hanlon, R. T. and J. B. Messenger. 1996. Cephalopod Behavior. Cambridge: Cambridge University Press, 1996.
Hanlon, R. T., M. J. Naud, J. W. Forsythe, K. Hall, A. C. Watson, and J. McKechnie. 2007. Adaptable night camouflage by cuttlefish. Am Nat 169: 543-551
Mathger, L. M., A. Barbosa, S. Miner, and R. T. Hanlon. 2006. Color blindness and contrast perception in cuttlefish (Sepia officinalis) determined by a visual sensorimotor assay. Vision Research 46(11): 1746-1753.
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Reproductive Behavior - Overview
Every winter thousands of Giant Cuttlefish (Sepia apama Gray) aggregate to spawn along a restricted area of rocky reef in northern Spencer Gulf, South Australia. It is the only known spawning aggregation of cuttlefish in the world and represents an exceptional cuttlefish mating system of high complexity. Females move around the area paying little heed to males, and males pursue females and compete for them. S. apama male tactics are diverse. Females attach their eggs individually to the underside of rocks, ledges, and caves in subtidal rocky reef habitats, and the eggs hatch in 3–5 months.
References
Hall, K. C. and R. T. Hanlon. 2002. Principal features of the mating system of a large spawning aggregation of the giant Australian cuttlefish Sepia apama (Mollusca : Cephalopoda). Mar Biol 140: 533-545
The University of Adelaide News Story: Cross-dressing cuttlefish to sex up tourism
Discover Magazine: Cuttlefish in Love
The Independent News Article: Cross-dressing cuttlefish is Casanova of the reef
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Male Behavior
Since the sex ratio on the spawning grounds ranges from 4-11 males/1 female, there is a high level of competition for females. Males on the spawning grounds are sexually dimorphic by size. Large males (mean 370 mm mantle length, ML) pair with and guard females (mean 250 mm ML) temporarily (pre- and postcopulation). Large lone males search for lone females or challenge consorts with agonistic displays. Large males engage in agonistic contests that pass through increasingly complex stages of display, culminating in a dramatic passing cloud display and sometimes physical contact and biting (see video of courtship behavior). Contests ended with one male swimming away. Small lone males guard females if there is no large male in the vicinity, or search for lone females with whom they mate without guarding. Small males (150–250 mm ML) may also use opportunities for extrapair copulations (EPCs). They use "open stealth" (overt sneak mating), "hidden stealth" (concealed sneak mating, e.g. under a rock) and female mimicry to achieve sneaker EPCs (see video of males mimicking females). Female mimicry leads to increased acceptance of mating with the female, and to immediate fertilization, as demonstrated by DNA fingerprinting.
References
Hall, K. C. and R. T. Hanlon. 2002. Principal features of the mating system of a large spawning aggregation of the giant Australian cuttlefish Sepia apama (Mollusca : Cephalopoda). Mar Biol 140: 533-545
Hanlon, R. T., M. J. Naud, P. W. Shaw, and J. N. Havenhand. 2005. Behavioural ecology - Transient sexual mimicry leads to fertilization. Nature 433: 212-212
Naud, M. J., R. T. Hanlon, K. C. Hall, P. W. Shaw, and J. N. Havenhand. 2004. Behavioural and genetic assessment of reproductive success in a spawning aggregation of the Australian giant cuttlefish, Sepia apama. Anim Behav 67: 1043-1050
Norman, M. R. 2000. Cephalopods, a world guide : Pacific Ocean, Indian Ocean, Red Sea, Atlantic Ocean, Caribbean, Arctic, Antarctic, ConchBooks, Hackenheim, Germany 318p
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Female Behavior
Females ultimately determine whether to accept a mating attempt. They reject a large percentage of mating attempts (up to 70%) and sometimes reject large consort males to mate with a small lone male. Females possess a sperm receptacle below the mouth and may use stored sperm from this receptacle or sperm from spermatangia placed in the buccal area to fertilize their eggs. Thus, females may accumulate sperm from consecutive matings and create considerable potential for sperm competition; it is not known if females exert cryptic sperm choice. Individual females can be extremely active on the spawning grounds: in a single day one female was documented to mate 17 times, with 8 different males, and lay 37 eggs in the course of 9 hours.
References
Hall, K. C. and R. T. Hanlon. 2002. Principal features of the mating system of a large spawning aggregation of the giant Australian cuttlefish Sepia apama (Mollusca : Cephalopoda). Mar Biol 140: 533-545
Hanlon R, T. and J. B. Messenger. 1996. Cephalopod Behavior. Cambridge: Cambridge University Press, 1996.
Naud, M. J., R. T. Hanlon, K. C. Hall, P. W. Shaw, and J. N. Havenhand. 2004. Behavioural and genetic assessment of reproductive success in a spawning aggregation of the Australian giant cuttlefish, Sepia apama. Anim Behav 67: 1043-1050
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Mating
Female Sepia apama alternate mating with egg-laying and often mate with multiple males between egg-laying events. Mating occurs in the head-to-head position during which the male flushes water over the buccal area before placing spermatophores there. Eggs are extruded singly and held amidst the female's arms - this appears as a bulge in her arms; at this time, sperm from the spermatangia in that area, or sperm released from the sperm receptacle, compete to fertilize the egg. Females then attach their eggs individually to the underside of rocks, ledges and reefs in subtidal rocky habitat.
Reference
Hall, K. C. and R. T. Hanlon. 2002. Principal features of the mating system of a large spawning aggregation of the giant Australian cuttlefish Sepia apama (Mollusca : Cephalopoda). Mar Biol 140: 533-545.
Naud, M. J., R. T. Hanlon, K. C. Hall, P. W. Shaw, and J. N. Havenhand. 2004. Behavioural and genetic assessment of reproductive success in a spawning aggregation of the Australian giant cuttlefish, Sepia apama. Anim Behav 67: 1043-1050.
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Life Cycle
Life History
Sepia apama have a life span of 1-2 years. Not much is known about their life history outside of the breeding season. Breeding takes place during the austral winter (May to September), when hundreds of thousands of animals aggregate in northern Spencer Gulf to mate and lay eggs. Recent research based on analysis of growth increments of cuttlebones suggests that Sepia apama has two alternative life cycles (for both sexes). The first involves rapid juvenile growth during the first summer after hatching, with maturity reached within 7–8 months. These individuals return to spawn in their first year as small individuals. The second life cycle involves much slower juvenile growth during the first summer, with maturity deferred until their second year, when they return to spawn as much larger individuals.
Reference
Hall, K. C., A. J. Fowler, and M. C. Geddes. 2007. Evidence for multiple year classes of the giant Australian cuttlefish Sepia apama in northern Spencer Gulf, South Australia. Rev Fish Biol Fish 17: 367-384.
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Evolution and Systematics
Systematics or Phylogenetics
Classification
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Order: Sepiida
Family: Sepiidae
Genus: Sepia
Species: Sepia apama
References
Gray, J. E. 1849. Catalogue of the Mollusca in the British Museum. Part I. Cephalopoda Antepedia. 164.
Lipinski, M. R., F. A. Naggs, and M. A. Roeleveld / N. A. Voss, M. Vecchione et al., eds. 1998. Nominal type specimens of Sepiidae in The Natural History Museum, London (BMNH). Systematics and Biogeography of Cephalopods. Smithsonian Contributions to Zoology, 586 (I-II): 157.
Lu, C. C. / N. A. Voss, M. Veccione, R. B. Toll and M. J. Sweeney, eds. 1998. A synopsis of Sepiidae in Australian waters (Sepioidea: Cephalopoda). Systematics and Biogeography of Cephalopods. Smithsonian Contributions to Zoology, 586 (I-II): 159-190.
Sweeney, M. J. and C. F. E. Roper / N. A. Voss, M. Vecchione, R. B. Toll and M. J. Sweeney, eds. 1998. Classification, type localities and type repositories of recent Cephalopoda. Systematics and Biogeography of Cephalopods. Smithsonian Contributions to Zoology, 586 (I-II): 561-599.
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Concepts and Synonymy
Amplisepia parysatis Iredale, 1954, Sepia palmata Owen, 1881
References
Iredale, T. 1954. Cuttle-fish "Bones" again. Australian Zoologist 1(1): 63-82.
Rudman, W. B. 1983. The Cephalopod Collections of the Australian Museum. Memoirs of the Museum of Victoria 44: 67-68.
Lu, C. C. / N. A. Voss, M. Veccione, R. B. Toll and M. J. Sweeney, eds. 1998. A synopsis of Sepiidae in Australian waters (Sepioidea: Cephalopoda). Systematics and Biogeography of Cephalopods. Smithsonian Contributions to Zoology, 586 (I-II): 159-190.
Owen, R. 1881. Descriptions of some new and rare Cephalopoda (Part II). Transactions of the Zoological Society of London 11(5): 131-170.
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Physiology and Cell Biology
Physiology
Vision
Cuttlefish have a uniquely shaped pupil comprising several slits in different orientations - some are W-shaped and others, like S. apama, have one horizontal slit that bends obliquely vertical. Light entering the eye is controlled by altering the shape of the entire eyeball. To focus on nearby objects a cuttlefish contracts a surrounding layer of the muscle, moving the lens forward, away from the retina and it focuses on distant objects by drawing the lens in. Cuttlefish are known to be color-blind, having only one visual pigment at 492nm. Despite being color-blind cuttlefish are very good at blending into colorful natural environments (at least in shallow depths of water).
References
Oceanworld Cuttlefish Fact Files
Marshall, N. J. and J. B. Messenger. 1996. Colour-blindcamouflage. Nature 382: 408–409
Mathger, L., A. Barbosa, S. Miner and R. T. Hanlon. 2006. Color blindness and contrast perception in cuttlefish (Sepia officinalis) determined by a visual sensorimotor assay. Vision Res. 46: 1746-1753.
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Physiology and Biochemistry
A recent energetics study found that Sepia apama are primarily diurnal and have a small home range (90-550 meters) over short recording periods. They are able to channel most of their energy directly into growth because they spend 95% of the day resting, suggesting bioenergetics more like that of an octopus than a squid. Very little time is spent foraging (3.7% during the day and 2.1% during the night), most of their time is spent resting and hiding in crevices from predators. The exception to this behavioral routine is the mass spawning aggregation, where cuttlefish are far more active during the days or weeks that they spend there.
References
Aitken, J. P., R. K. O'Dor and G. D. Jackson. 2005. The secret life of the giant Australian cuttlefish Sepia apama (Cephalopoda): Behaviour and energetics in nature revealed through radio acoustic positioning and telemetry (RAPT). J Exp Mar Biol Ecol 320: 77-91
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Molecular Biology and Genetics
Molecular Biology
Molecular Biology and Genetics
Several microsatellite markers have been produced for the assessment of genetic diversity and paternity analyses in Sepia apama (Genbank accession numbers EF368216-224, AY617047, AY617024, AY616977, AY616950, AF533071-8, AY294335-353). Recent genetic evidence (using allozyme electrophoresis, microsatellites and mitochondrial nucleotide sequences) has revealed three major genetically distinct groups of Sepia apama. It has been recommended that each of these groups be managed separately to preserve genetic diversity. Initial paternity analysis of the complex mating system of the giant cuttlefish indicates that this mating system has a high level of multiple mating that results in multiple paternity; males of any size or status can obtain successful fertilizations. Genetic analysis has also shown that the two female sperm storage areas (the short-term spermatangia - in the buccal area- and the long-term sperm receptacle - beneath the beak) both can contain sperm from multiple males. Of the two storage areas, sperm from the spermatangia (recent matings within a few hours or days) is used more frequently to fertilize eggs than sperm from the receptacle.
References
Kassahn, K. S., S. C. Donnellan, A. J. Fowler, K. C. Hall, M. Adams, and P. W. Shaw. 2003. Molecular and morphological analyses of the cuttlefish Sepia apama indicate a complex population structure. Mar Biol 143: 947-962.
Naud, M. J., P. W. Shaw, R. T. Hanlon, and J. N. Havenhand. 2005. Evidence for biased use of sperm sources in wild female giant cuttlefish (Sepia apama). Proc R Soc Lond Ser B-Biol Sci 272: 1047-1051.
Naud, M. J., R. T. Hanlon, K. C. Hall, P. W. Shaw, and J. N. Havenhand. 2004. Behavioural and genetic assessment of reproductive success in a spawning aggregation of the Australian giant cuttlefish, Sepia apama. Anim Behav 67: 1043-1050.
Shaw, P. W. 2003. Polymorphic microsatellite DNA markers for the assessment of genetic diversity and paternity testing in the giant cuttlefish, Sepia apama (Cephalopoda). Conserv Genet 4: 533-535.
Wheaton, L., S. C. Donnellan, M. C. De, M. G. Gardner, and B. M. Gillanders. 2007. Isolation of additional polymorphic tri- and tetranucleotide microsatellite loci for the giant Australian cuttlefish (Sepia apama). Mol Ecol Notes 7: 893-895.
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Conservation
Conservation and Fisheries
Prior to mid 1990s, the spawning aggregation at Whyalla, in northern Spencer Gulf, was fished at sustainable levels for snapper bait. However, in the mid 1990’s, fishers actively targeted cuttlefish, and large numbers of the breeding aggregation were removed from the system. Since the lifecycle of Sepia apama is very short (1-2 yrs), if a cohort of breeders is fished out, the following generation is likely to be severely impacted. To avoid long-term population decline, even local extinction, a renewable moratorium preventing fishing was introduced in 1999. In subsequent years, the cuttlefish numbers increased again, and ecotourism in the area began to thrive, as it does at this writing (2007). The most recent survey, done in 2005, showed a slight decline in abundance since the estimates made in 1998 and 2001. This survey indicates that declines may be the result of multiple factors - not solely over-exploitation from targeted human fishing. More precise molecular data on population genetics will help determine the composition of the hundreds of thousands of cuttlefish that constitute this extremely unique mass spawning aggregation. This is a clear case of ecotourism aiding conservation.
References
The University of Adelaide Cuttlefish Project
Steer, M. A. and K. C. Hall. 2005. Estimated abundance and biomass of the unique spawning aggregation of the giant Australian cuttlefish (Sepia apama) in northern Spencer Gulf, South Australia. Report to Coastal Protection Branch, Department for Environment and Heritage, South Australia. South Australian Research and Development Institute (Aquatic Sciences), Adelaide, RD 05/0012-1.
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Threats
How to Grow
Dr. Karina Hall (2002) reared the eggs to juvenile adult stage but growth was very slow compared to that of other cuttlefish such as Sepia officinalis. A great deal more aquaculture effort on a larger scale is needed to assess the convenience of culturing this species in the laboratory or for providing a food source. The largest problem is that cuttlefish are carnivores that strongly prefer live food such as crustaceans and fish.
References
Hall, K. C. 2000. Cuttlefish mysteries. Southern Fisheries 7
Forsythe, J. W., R. H. DeRusha, and R. T. Hanlon. 1994. Growth, reproduction and life span of Sepia officinalis (Cephalopoda: Mollusca) cultured through seven consecutive generations. J. Zool. 233: 175–192.
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Relevance to Humans and Ecosystems
Benefits
Uses
Since ca. 1999, this species has been an important ecotourism attraction to the city of Whyalla, South Australia. Divers come from around the world to see this amazing underwater spectacle of thousands of active cuttlefish on the spawning aggregation. This spectacle has grown in its popularity and continues to be featured in international television productions around the world. There is even a bus tour that features the giant cuttlefish (see image below). Outside of the protected spawning areas, this cuttlefish supports a small scale bait fishery, and some animals are eaten by humans. Importantly, this highly unique spawning aggregation has become a biological model for mating systems (see many scientific journal papers published on this since 1999).

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Wikipedia
Sepia apama
Sepia apama, also known as the Australian giant cuttlefish, is the world's largest cuttlefish species, growing to 50 cm in mantle length and over 10.5 kg (23 lb) in weight.[2] Using cells known as chromatophores, the cuttlefish can put on spectacular displays, changing colour in an instant.
S. apama is native to the southern coast of Australia, from Brisbane in Queensland to Shark Bay in Western Australia. It occurs on rocky reefs, seagrass beds, and sand and mud seafloor to a depth of 100 m.[3]
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Life cycle and reproduction [edit]
Sepia apama live from two to three years. Breeding takes place with the onset of the southern winter. Males abandon their normal cryptic colouring and set out to dazzle the females by adopting rapidly changing bright colours and striking patterns. Females are polyandrous, and collaborative research indicates the tendency for females to reproduce using male genetic material deposited in spermatangia more favorably than in sperm receptacles directly. Females then attach their eggs to the underside of rocks in caves or crevises where they will hatch within three to five months. Sepia apama are semelparous and death follows shortly after a single mating cycle and laying of eggs that will spawn the next generation.[4] Sepia apama have poor anaerobic capability compared to most aquatic invertebrates and a lack of food leads to catabolism. Stomach content analysis indicates fasting during the breeding season and as Sepia apama can catabolise no more than 50% of their body weight they slowly lose physical condition as the season progresses and eventually die. Throughout their range, Sepia apama breed in pairs or small groups, laying eggs in suitable caves or rock crevises. Loose spawning aggregations can form but rarely exceed 10 animals in any one location,[5] with one known exception: hundreds of thousands aggregate off Point Lowly in the Upper Spencer Gulf. While surveys suggest that juveniles leave the spawning ground after hatching, nothing is known of their subsequent movement or lifestyle strategies as a juvenile.
Physiology and biochemistry [edit]
Genetic studies have shown that there is little if any interbreeding between Sepia apama populations. While there is some genetic divergence, the various populations are not considered taxonomically distinct and are commonly referred to by their location, e.g. Sepia apama upper Spencer Gulf population.[4] The upper Spencer Gulf population is unique in that a permanent salinity gradient in the Spencer Gulf may physiologically exclude other populations from the zone occupied by the upper Spencer Gulf population.[4] Research suggests that the upper Spencer Gulf population may in fact be a separate species as it does show some hallmarks, such as genetic separation, differences in morphology and different patterns of sexual dimorphism to adjacent populations.[5][6]
Sepia apama is a neritic demersal species. They are carnivorous, opportunistic and voracious predators who feed predominantly on crustaceans and fish.[5] Using neurally controlled cells known as chromatophore organs (red to yellow), iridophores (iridescent: spans the entire visible spectrum from blue to near-IR) and leucophores (white), the cuttlefish can put on spectacular displays, changing colour and patterns in a fraction of a second. Located in three layers under the skin, leucophores make up the bottom layer, with chromatophores the outermost. By selective blocking, the three layers work together to produce polarised patterns. Unlike those in most animals, cuttlefish iridophores are physiologically active; they can change their reflectivity and the degree of polarization can also be controlled. Cuttlefish are colourblind, however the photoreceptors of cuttlefish eyes are arranged in a way which gives them the ability to see the linear polarization of light. While the Mantis shrimp is the only known creature to have true polarization vision, it is believed that cephalopods may also.[7] Because the optic lobes of cuttlefish are larger than any other region of the brain and their skin produces polarized reflective patterns, it has been postulated that they may communicate through this visual system.[8] By raising elaborate papillae on their skin, S. apama can change the shape and the texture of their skin to imitate rock, sand or seaweed.[9]
A recent energetics study found that Sepia apama are primarily diurnal and have a small home range (90–550 metres) over short recording periods while travelling large distances to breed. They are able to channel most of their energy directly into growth because they spend 95% of the day resting, suggesting bioenergetics more like that of an octopus than a squid. Very little time is spent foraging (3.7% during the day and 2.1% during the night), most of their time is spent resting and hiding in crevices from predators. The exception to this behavioral routine is the mass spawning aggregation, where cuttlefish are far more active during the days or weeks that they spend there.[10][11]
Role in ecosystem [edit]
The Australian giant cuttlefish is eaten by Indo-Pacific bottlenose dolphins, which have been observed (in South Australia's Spencer Gulf) to have developed a technique for removing the ink and cuttlebone from a cuttlefish before eating it.[12]
Upper Spencer Gulf population [edit]
Discovered by divers in the late 1990s, the upper Spencer Gulf population is the world's only known mass cuttlefish spawning aggregation with hundreds of thousands of Sepia apama congregating on sub-tidal reefs around Point Lowly near Whyalla between May and August. While outside of the breeding season, the sex ratio is 1 to 1, Spencer Gulf males outnumber females by up to 11 to 1 in the spawning aggregation. It is unclear if this is due to fewer females taking part or to males breeding for a longer period of time than females. With densities of one cuttlefish per square metre, covering approximately 61 hectares (150 acres), the sheer number of Sepia apama makes this breeding aggregation unique in the world. As the cuttlefish are oblivious to divers while spawning, they are now a major regional tourist attraction for divers from around the world.[5] Professor Roger Hanlon of the Woods Hole Oceanographic Institution has called the breeding aggregation "the premier marine attraction on the planet."[6]
The Sepia apama upper Spencer Gulf population displays two alternative life cycles in both sexes (growth pattern polymorphism). The first involves rapid growth with maturity reached in seven to eight months with small adults returning to spawn in the first year. The second involves slow growth with maturity reached in two years with large adults returning to spawn in the second year.[4] The upper Spencer Gulf population displays reproductive behaviours unique to this population, possibly as a result of the high spawning densities. Large males defend females and egg laying sites while small males mimic female colouring and form in order to gain access to the females being protected by the dominant males which are extremely territorial. Male genetic material is deposited in sperm receptacles directly. The females, who potentially lay hundreds of eggs, extracts one egg at a time and fertilises it by passing it over the sperm receptacle before attaching it to the underside of a rock at depths of 2 metres (6.6 ft) to 5 metres (16 ft).[5]
Prior to the mid-1990s, the population was fished for snapper bait with annual catches of around four tonnes (approx 4,000 cuttlefish). During the 1995 and 1996 spawning seasons, commercial fishing of the spawning grounds harvested around 200 tonnes annually. Over exploitation was recognised after 245 tonnes was harvested in 1997, leading to 50% of the grounds being closed to commercial fishing in 1998. Despite half of the grounds being closed, commercial fishers took 109 tonnes in 1998 (approximately half of the estimated biomass) before dropping to 3.7 tonnes in 1999. The catch data for 2000 to 2005 has not been released for confidentiality reasons.[13] Surveys indicated that the cuttlefish biomass remained stable from 1998 to 2001. A further survey in 2005 revealed a 34% decrease in biomass since 2001 that was attributed to natural variability and illegal fishing during the peak spawning period.[13] The closure was subsequently expanded to the entire spawning grounds and anecdotal observations suggested increased numbers in 2006 and 2007, however, a new survey in 2008 found the biomass had decreased a further 17%.[4][5]
In 2011, it was estimated that only 33% of the 2010 population had returned to breed, less than 80,000 cuttlefish. Beginning in May, the cuttlefish leave deep water and migrate along coastal reefs to reach their spawning grounds. Local fishermen claim that a small "finger of land" near Point Lowly extends outside the exclusion zone and that commercial fishers have been targeting the area, intercepting the Sepia apama before they can reach the spawning grounds. Being semelparous breeders, ecologist Bronwyn Gillanders believes the cuttlefish to be in danger, stating that it is hard to determine whether this a natural phenomenon or something else and that the cause requires more research.[14]
In 2012, the number of cuttlefish that returned to the spawning ground again dropped dramatically with numbers as low as 6,000 estimated. A state government working group had been unable to determine the cause of the 2011 decline and have now recommended investigating broader ecological factors. Tour guide Tony Bramley, who had been taking divers to view the spawning grounds since they were discovered stated "It's heartbreaking, when you look at what's left...[once] there were so many animals you couldn't land on the bottom, you had to push them aside."[15] The Conservation Council of SA, which believes the population to be a separate species, has warned that the Spencer Gulf cuttlefish face extinction within two or three years if nothing is done to protect them. The state government working group had recommended an immediate ban on fishing for the cuttlefish, however, this was rejected by State Cabinet on 3 September with Fisheries Minister Gail Gago stating; "There is no strong evidence to suggest that fishing is impacting on the giant cuttlefish, therefore, further closures would be ineffective."[6]
Effect of local industrialisation [edit]
In 1984, before the spawning grounds were discovered, Santos built a hydrocarbon processing plant at adjoining Port Bonython. There is some concern over the possible impact of the plant on the cuttlefish population and there have been two major spills at the plant.[16][17] Santos denies that the recent spill spread off-site, but the SA EPA said hydrocarbons had migrated through the rock strata beyond the plant and the barrier trench built by Santos. Santos now provides funding for cuttlefish research.[18]
BHP Billiton has plans to build a desalination plant at Point Lowly to supply fresh water to Roxby Downs copper and uranium mine. The plant, located within 200 metres (660 ft) of the breeding grounds, would release around 120 megalitres (32,000,000 US gal) of brine (46–60 ppt) into the area each day. As cuttlefish embryos die off as salinity levels rise (optimal range 28–38 ppt, 100% mortality at 50 ppt), there has been considerable public opposition to the proposed plant because of the expected environmental impact.[18][19][20]
Due to its proximity to the ore deposits of the Middleback Ranges, several mining companies have indicated they want to build petrochemical and diesel refining facilities at Point Lowly. A second jetty for the loading of iron ore, and possibly copper and uranium, is also planned. The Cuttlefish Coast Coalition has been formed to fight these proposed developments.[18]
As a result of the above threats to the population, in 2010 an application was made to list Sepia apama in the list of threatened species. On February 2, 2011, the Threatened Species Scientific Committee ruled the species was not eligible for listing as the affected population was not taxonomically distinct from the rest of Sepia apama for the purposes of the Act.[20]
See also [edit]
References [edit]
- ^ Barratt, I. & Allcock, L. 2012. Sepia apama. In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. <www.iucnredlist.org>. Downloaded on 27 October 2012.
- ^ Reid, A., P. Jereb, & C.F.E. Roper 2005. Family Sepiidae. In: P. Jereb & C.F.E. Roper, eds. Cephalopods of the world. An annotated and illustrated catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes. No. 4, Vol. 1. Rome, FAO. pp. 57–152.
- ^ Norman, M.D. 2000. Cephalopods: A World Guide. ConchBooks.
- ^ a b c d e Amendment to the list of Threatened Population under the Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act). Threatened Species Scientific Committee.
- ^ a b c d e f Natural History of the Giant Australian Cuttlefish BHP Billiton
- ^ a b c Clare Peddie, Heather Kennett The cuttle scuttle The Advertiser September 29, 2012 Pg 67
- ^ Silvery fish reflect polarised light; they reflect the same amount of light in the same direction as the light they are viewed against, making them almost invisible in water. If the polarisation is reduced, the fish become easily visible. Cuttlefish will ignore fish with reduced polarisation and preferentially attack fish emitting polarised light.[1]
- ^ Mäthger et al (April 21, 2009). "Do cephalopods communicate using polarized light reflections from their skin?". The Journal of Experimental Biology 212, 2133-2140. Retrieved 4 November 2011.
- ^ Alison King The Colourful World of Cephalopods - Cephalopod body patterning II. The Cephalopod Page.
- ^ Hanlon, R.T. 2008. Australian Giant Cuttlefish - Physiology and Biochemistry. Encyclopedia of Life.
- ^ Aitken, J.P., R.K. O'Dor & G.D. Jackson. 2005. The secret life of the giant Australian cuttlefish Sepia apama (Cephalopoda): Behaviour and energetics in nature revealed through radio acoustic positioning and telemetry (RAPT). Journal of Experimental Marine Biology and Ecology 320: 77–91.
- ^ Catch cuttlefish, drain off the ink, then fillet. Serves five (dolphins): Scientists stunned by mammals' elaborate culinary preparations
- ^ a b MA Steer and KC Hall Estimated Abundance and Biomass of the Unique Spawning Aggregation of the Giant Australian Cuttlefish (Sepia apama) in Northern Spencer Gulf, South Australia. PIRSA October 2005
- ^ Mystery of the Missing Cuttlefish. The Advertiser, July 24, 2011. p. 7.
- ^ Drastic loss in giant colony. The Advertiser, September 25, 2012. p. 24.
- ^ http://www.amsa.gov.au/marine_environment_protection/major_oil_spills_in_australia/Era/index.asp
- ^ http://www.abc.net.au/news/2010-03-22/epa-fears-bigger-santos-oil-spill/374516
- ^ a b c Cuttlefish chaos at Whyalla. Archived from Dive Pacific Issue No. 116, Feb/Mar 2010.
- ^ Desalination and South Australia's Gulfs ecosystems. Fishers For Conservation Inc.
- ^ a b Conservation Assessment. Advice to the Minister for Sustainability, Environment, Water, Population and Communities from the Threatened Species Scientific Committee (the Committee) on Amendment to the list of Threatened Population under the Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act) February 2, 2011.
- Kassahn, K.S., S.C. Donnellan, A.J. Fowler, K.C. Hall, M. Adams & P.W. Shaw 2003. Molecular and morphological analyses of the cuttlefish Sepia apama indicate a complex population structure. Marine Biology 143(5): 947–962. doi:10.1007/s00227-003-1141-5
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