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. 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.
Life cycle and reproduction
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. 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 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
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. 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. 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.
Sepia apama is a neritic demersal species. They are carnivorous, opportunistic and voracious predators who feed predominantly on crustaceans and fish. 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. 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. 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.
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.
Role in ecosystem
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.
Upper Spencer Gulf population
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. Professor Roger Hanlon of the Woods Hole Oceanographic Institution has called the breeding aggregation "the premier marine attraction on the planet."
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. The upper Spencer Gulf population displays reproductive behaviours unique to this popuation, 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).
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. 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. 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%.
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.
In 2012, the number of cuttlefish that returned to the spawning ground again dropped dramaticaly 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." 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."
Effect of local industrialisation
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. 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.
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 USgal) 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.
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.
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.
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