28
proven that an enriched environment, more space, and better nutritional state may
keep the octopus confined. Still, some of them just want to leave. In the longer term,
octopuses may make onshore–offshore migrations, perhaps for reproduction or to
access a more appropriate food supply, but this may be irrelevant to their culture
unless light level affects their reproduction.
2.5.3 Semelparity
Coastal cephalopods typically mature in about a year (Boyle and Boletzky 1996)
and although many are semelparous, Rocha et al. (2001) have identified five different cephalopod reproductive strategies along a gradient in environmental stability,
including four varieties of multiple spawning, three of which are nonterminal. Recognizing this complexity in cephalopod spawning behaviours has blurred the sharp
distinction between semelparity and iteroparity. Cuttlefish lay individual or small
groups of eggs and sequester them in protected spaces. They spawn intermittently
over a period of several months. The squid deposit eggs in large egg masses, some
seasonally in mass synchronized spawning events, and are typified by terminal
spawning although there are some examples of nonterminal spawning (Rocha et al.
2001). According to Rocha et al. (2001), octopuses span the continuum between
semelparity and iteroparity: Many shallow-water octopuses are small-egged species
laying eggs in festoons or small clusters of a single spawning event, while deepwater octopuses such as Graneledone sp. but also some shallow-water octopuses
(Anderson and Wood 2012) are large-egged species depositing eggs singly, perhaps
over a protracted period. E� megalocyathus is an intermediate example (Ortiz 2006).
In either case, females brood eggs over a long period and die after hatching of the
young.
2.5.4 Mating and Spawning
Male cuttlefish court females, using multiple mating strategies (Norman et al.
1999), and deceive other males via female mimicry (Norman et al. 1999; Brown
et al. 2012). Mass spawning is seen in the giant Australian cuttlefish, S� apama
(Hall and Hanlon 2002). During these mass spawnings, both sexes have multiple
mates; males defended females but not egg-laying sites, and males attempted takeovers of paired females through agonistic displays and attention to opportunities for
extra-pair copulations via stealth or female mimicry (Hall and Hanlon 2002). Many
aspects of this interaction, but not mass spawning, are typical of Sepiida.
S� officinalis males present a striped body pattern (intense zebra display) to other
males; larger and darker males are dominant in such interactions (Boal 1997). However, females show consistent preference not for the larger or darker males, but for
the most recently mated males and those showing fewer zebra displays (Boal 1997).
She suggested that females used chemical cues rather than visual ones in assessing
J. Mather and D. Scheel
proven that an enriched environment, more space, and better nutritional state may
keep the octopus confined. Still, some of them just want to leave. In the longer term,
octopuses may make onshore–offshore migrations, perhaps for reproduction or to
access a more appropriate food supply, but this may be irrelevant to their culture
unless light level affects their reproduction.
2.5.3 Semelparity
Coastal cephalopods typically mature in about a year (Boyle and Boletzky 1996)
and although many are semelparous, Rocha et al. (2001) have identified five different cephalopod reproductive strategies along a gradient in environmental stability,
including four varieties of multiple spawning, three of which are nonterminal. Recognizing this complexity in cephalopod spawning behaviours has blurred the sharp
distinction between semelparity and iteroparity. Cuttlefish lay individual or small
groups of eggs and sequester them in protected spaces. They spawn intermittently
over a period of several months. The squid deposit eggs in large egg masses, some
seasonally in mass synchronized spawning events, and are typified by terminal
spawning although there are some examples of nonterminal spawning (Rocha et al.
2001). According to Rocha et al. (2001), octopuses span the continuum between
semelparity and iteroparity: Many shallow-water octopuses are small-egged species
laying eggs in festoons or small clusters of a single spawning event, while deepwater octopuses such as Graneledone sp. but also some shallow-water octopuses
(Anderson and Wood 2012) are large-egged species depositing eggs singly, perhaps
over a protracted period. E� megalocyathus is an intermediate example (Ortiz 2006).
In either case, females brood eggs over a long period and die after hatching of the
young.
2.5.4 Mating and Spawning
Male cuttlefish court females, using multiple mating strategies (Norman et al.
1999), and deceive other males via female mimicry (Norman et al. 1999; Brown
et al. 2012). Mass spawning is seen in the giant Australian cuttlefish, S� apama
(Hall and Hanlon 2002). During these mass spawnings, both sexes have multiple
mates; males defended females but not egg-laying sites, and males attempted takeovers of paired females through agonistic displays and attention to opportunities for
extra-pair copulations via stealth or female mimicry (Hall and Hanlon 2002). Many
aspects of this interaction, but not mass spawning, are typical of Sepiida.
S� officinalis males present a striped body pattern (intense zebra display) to other
males; larger and darker males are dominant in such interactions (Boal 1997). However, females show consistent preference not for the larger or darker males, but for
the most recently mated males and those showing fewer zebra displays (Boal 1997).
She suggested that females used chemical cues rather than visual ones in assessing
J. Mather and D. Scheel