The behavioural ecology of Sepiolidae (Cephalopoda: Sepiolida): a review

Figures & data

Figure 1. Sepiolid anatomy, using Sepiola atlantica as an example. A, lateral view; B, oral view. Abbreviations: chr —chromatophores; fun —funnel; LI (♂ hec) —left arm I (in male individuals: hectocotylus); LII —left arm II; LIII —left arm III; LIV —left arm IV; man —mantle; RI —right arm I; RII —right arm II; RIII —right arm III; RIV —right arm IV; suc —sucker; ten —tentacle. The 4 arm pairs consist of one arm from either side, e.g. LI and RI form the arm pair I, LII and RII form arm pair II, and so forth. Credits: A–B, Photos taken by Bas van der Sanden.

Figure 2. Hunting behaviour in sepiolids. A, B, An individual of Eumandya parva approaching its prey (here: mysid shrimp) with its arms forming an elongated cone; C–F, Tentacular strike to capture the prey; G, H, Retraction of tentacles to pull the prey towards the mouth. Credits: A–H, Reprinted from Drerup et al. (2020), with permission from Elsevier.

Table 1. Literature dedicated to different ecological aspects of sepiolids. Taxonomic authorities are given only for species not mentioned in the text.

Ecological aspect	Subfamily	Species	References
Feeding and hunting behaviour	Sepiolinae	Adinaefiola ligulata (Naef, 1912) Eumandya parva	Drerup et al. (2020)
		Euprymna berryi Sasaki, 1929	Choe (1966)
		Euprymna brenneri Sanchez et al., 2019	Sanchez et al. (2019)
		Euprymna hyllebergi	Nabhitabhata et al. (2005)
		Euprymna scolopes	Moynihan (1983); Shears (1988); Hanlon et al. (1997)
		Sepietta neglecta Naef, 1916	Boletzky et al. (1971)
		Sepietta obscura Naef, 1916	Boletzky et al. (1971)
		Sepietta oweniana	Bergström (1985); Orsi Relini and Massi (1988),
		Sepiola affinis	Mauris (1989)
		Sepiola atlantica	Yau and Boyle (1996); Rodrigues et al. (2011c)
		Sepiola robusta	Boletzky et al. (1971); Boletzky (1983)
		Sepiola rondeleti Leach, 1817	Boletzky et al. (1971)
			Boletzky et al. (1971)
	Rossiinae	Rossia macrosoma	Boletzky and Boletzky (1973); Drerup et al. (2021)
		Rossia pacifica	Brocco (1971); Summers and Colvin (1989)
Anti-predator behaviour	Sepiolinae	Eumandya parva	Drerup et al. (2020)
		Euprymna hyllebergi	Nabhitabhata et al. (2005)
		Euprymna scolopes	Singley (1982); Moynihan (1983); Singley (1983); Shears (1988); Anderson and Mather (1996); Jones and Nishiguchi (2004); Stabb (2006); Stabb and Millikan (2009); Seehafer et al. (2018)
		Euprymna tasmanica	Norman and Lu (1997)
	Rossiinae	Rossia macrosoma	Drerup et al. (2021)
		Rossia pacifica	Shimek (1983)
	Heteroteuthinae	Heteroteuthis dispar	Dilly and Herring (1978)
Burying behaviour	Sepiolinae	Eumandya parva	Drerup et al. (2020)
		Euprymna scolopes	Moynihan (1983); Anderson (1997); Anderson et al. (2002)
		Euprymna hyllebergi	Nabhitabhata et al. (2005)
		Sepietta neglecta	Boletzky and Boletzky (1970)
		Sepietta obscura	Boletzky and Boletzky (1970)
		Sepiola atlantica	Rodrigues et al. (2010)
		Sepiola affinis	Boletzky and Boletzky (1970)
		Sepiola ligulata	Boletzky and Boletzky (1970)
		Sepiola robusta	Boletzky and Boletzky (1970); Boletzky (1983)
		Sepiola rondeletii	Boletzky and Boletzky (1970)
		Sepiola sp.	Drerup (2019)
	Rossiinae	Rossia macrosoma	Boletzky and Boletzky (1973); Drerup et al. (2021)
		Rossia pacifica	Anderson et al. (2004)

Figure 3. Anti-predator behaviour in sepiolids. A, B, The stout bobtail squid Rossia macrosoma occupies habitats with both soft and hard substrate and can either bury itself in soft sediment or shelter in rock crevices to avoid predation; C, Individual of Eumandya parva ejecting a stretch of ink, a so-called ‘ink rope’, and holding on to it. The picture of a captive individual was taken with an infrared-sensitive camera, however this behaviour has also been observed in the wild (Drerup et al. 2020); D, Individual of E. parva wearing a layer of sediment (‘sand coat’) covering its dorsal mantle; E, Counterillumination behaviour of sepiolids. Left: Species devoid of a light organ cast a shadow below their body. Right: Counterillumination by matching ambient light in species with light organs. Credits: A, B, Photos taken by Mark Skea and reprinted from Drerup et al. (2021) (CC BY); C, D, Reprinted from Drerup et al. (2020), with permission from Elsevier; E, Redrawn from Stabb (2006), with permission from John Wiley and Sons.

Figure 4. Burying behaviour in sepiolids. A, An individual of Sepiola intermedia shortly before resting on the sediment; B, Settling/resting posture before the start of burying; C, D, Phase 1 of the burying behaviour. After the formation of a depression in the sediment by tilting the body slightly forwards and ejecting a gentle forward-directed water jet, alternating backward- and forward-directed water jets are ejected to cover nearly the whole body with sediment; E, Phase 2 of the burying behaviour. A series of arm sweeps is conducted to gather sediment from its circular vicinity to cover the remaining body parts; C–E, During the burying procedure, the animal shifts from a dark to a pale body colouration; F, Fully buried individual. Credits: A–F, Photos taken by Christian Drerup.

Table 2. Literature dedicated to reproductive biology and egg characteristics of sepiolids. Taxonomic authorities are given only for species not mentioned in the text or Table 1.

Subfamily	Species	References
Sepiolinae	Eumandya parva	Sanchez et al. (2019)^d; Drerup et al. (2020)^a,b,c,d
	Euprymna berry	Choe (1966)^d
	Euprymna brenneri	Sanchez et al. (2019)^d
	Euprymna hyllebergi	Nabhitabhata et al. (2005)^a,b,c,d
	Euprymna scolopes	Arnold et al. (1972)^d; Moynihan (1983)^b; Singley (1983)^b; Hanlon et al. (1997)^b,c,d; Claes and Dunlap (2000)*
	Euprymna tasmanica	Norman and Lu (1997)^c,d; Steer et al. (2004)^d; Franklin et al. (2012)^e; Squires et al. (2012)^e; Squires et al. (2013)^a,b,c,d,e; Franklin et al. (2014)^e; Squires et al. (2014)^e; Squires et al. (2015)^e; Franklin and Stuart-Fox (2017)^e
	Rondeletiola minor	Salman and Katagan (1996)*; Czudaj et al. (2013)*; Önsoy et al. (2013)*; Akalin and Salman (2018)*
	Sepietta neglecta	Boletzky et al. (1971)^c,d; Lefkaditou and Kaspiris (1998)*; Deickert (2009)*; Akalin and Salman (2018)*
	Sepietta obscura	Boletzky et al. (1971)^c,d; Deickert (2009)*; Bello and Deickert (2021)^a,b
	Sepietta oweniana	Thorson (1946)*; Mangold-Wirz (1963)*; Bergstrom and Summers (1983)^b,c,d; Salman (1998)*; Bello and Deickert (2003)^c,d; Deickert and Bello (2005)^d; Cuccu et al. (2010)^d; Czudaj et al. (2012)*; Akalin and Salman (2018)*
	Sepiola spp.	Salman and Önsoy (2004)*
	Sepiola affinis	Mangold-Wirz (1963)^b; Gabel-Deickert (1995)^c,d; Bello and Deickert (2021)^a,b
	Sepiola atlantica	Levy (1912)^b; Yau and Boyle (1996)*; Rodrigues et al. (2009)^a,b; Jones and Richardson (2010)^b,c,d; Rodrigues et al. (2011b)^c,d; Rodrigues et al. (2011c)^c,d; Jones and Richardson (2012)*; Rodrigues et al. (2012)*; Goud et al. (2019)^b
	Sepiola intermedia Naef, 1912	Salman and Önsoy (2004)*; Salman (2014)*; Akalin and Salman (2018)*; Bello and Deickert (2021)^a,b
	Sepiola ligulata	Boletzky et al. (1971)^c,d
	Sepiola robusta	Boletzky et al. (1971)^c,d; Boletzky (1983)^b,c,d; Salman and Önsoy (2004)*; Akalin and Salman (2018)*
	Sepiola rondeletii	Racovitza (1894)^b; Boletzky et al. (1971) ^c,d; Akalin and Salman (2018)*
	Sepiola steenstrupiana Levy, 1912	Salman and Önsoy (2004)*; Akalin and Salman (2018)*
Rossiinae	Neorossia caroli	Cuccu et al. (2007)^d; Laptikhovsky et al. (2008)*; Salman (2010)*; Cuccu et al. (2011)*
	Rossia sp.	Aldrich and Lu (1968)^d
	Rossia macrosoma	Racovitza (1894)^b; Mangold-Wirz (1963)^b; Boletzky and Boletzky (1973)^d; Rosa et al. (2006)*; Laptikhovsky et al. (2008)*; Salman and Önsoy (2010)*; Akalin et al. (2016)^b; Drerup et al. (2021)^b,c,d
	Rossia moelleri	Laptikhovsky et al. (2008)*; Hoving et al. (2009)*
	Rossia mollicella Sasaki, 1920	Okutani and Sasaki (2007)^d
	Rossia pacifica	Brocco (1971)^b,c; Summers and Colvin (1989)^d; Anderson and Shimek (1994)^d; Mangold et al. (1998)^d; Laptikhovsky et al. (2008)*
	Semirossia patagonica (E. A. Smith, 1881)	Önsoy et al. (2008)*
Heteroteuthinae	Heteroteuthis dispar	Orsi Relini (1995)*; Hoving et al. (2008)*
	Stoloteuthis leucoptera	Villanueva and Sánchez (1993)*

^{a, b, c, d, e} Information on ^a (potential) courtship behaviour, ^b mating behaviour, ^c spawning behaviour, ^d eggs and ^e other reproductive behaviour. References marked with an asterisk (*) are not dedicated to a particular reproductive behaviour but provide useful information on the reproductive biology of the corresponding species.

Figure 5. Mating in sepiolids. A–C, Mating in sepioline species, using pairs of Sepiola atlantica as an example. Copulation usually occurs: A, B on the sediment, but occasionally C, pairs are also observed slowly swimming; D–G, Mating in rossiine sepiolids, here shown with individuals of Rossia macrosoma; D, Mating of two individuals; E–G, Due to the lack of a bursa copulatrix in female rossiine sepiolids, males implant their spermatophores (E) on the left ventral side close to the oviduct opening or (F–G) on the head or outer surface of the female’s mantle; G, Close-up of the dashed area in F. Credits: A, Photo taken by Mirjam van der Sanden. B, C, Photos taken by Bas van der Sanden. D–G, Photos taken by Mark Skea and reprinted from Drerup et al. (2021) (CC BY).

Figure 6. Spawning behaviour in sepiolids, here displayed by an individual of Rossia macrosoma. A–C, Spawning sequence. A, Female resting in front of an older egg batch after attaching one of her eggs to the old batch (marked with an arrow). B, The female displays her spawning behaviour by clinging to the substrate and spawning another egg. C, After successfully deploying one egg (marked with an arrow), the female rests for a moment then continues spawning individual eggs as shown above. D, Batch of freshly spawned eggs attached to a batch of older eggs, with some already hatched. Credits: A–D, Photos taken by Mark Skea and reprinted from Drerup et al. (2021) (CC BY).

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