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Growth and Mating of Southern African Lycoteuthis Lorigera (Steenstrup, 1875) (Cephalopoda; Lycoteuthidae)

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Growth and Mating of Southern African Lycoteuthis Lorigera (Steenstrup, 1875) (Cephalopoda; Lycoteuthidae) Rev Fish Biol Fisheries (2007) 17:259–270 DOI 10.1007/s11160-006-9031-9 ORIGINAL PAPER Growth and mating of southern African Lycoteuthis lorigera (Steenstrup, 1875) (Cephalopoda; Lycoteuthidae) H. J. T. Hoving Æ M. R. Lipinski Æ M. A. C. Roeleveld Æ M. D. Durholtz Received: 10 February 2006 / Accepted: 14 November 2006 / Published online: 23 December 2006 Ó Springer Science+Business Media B.V. 2006 Abstract Lycoteuthis lorigera is an oceanic males and females of equivalent ages. The rela- squid that is abundant in the Benguela system. tionship between ML and age for both sexes was Little is known about the biology of this squid best described by an exponential growth curve, except that it is eaten in large numbers by probably because no early life stages were aged in numerous oceanic predators and that males grow this study. Only one mature male (ML 160 mm) to larger size than females, which is unique for was aged, and preliminary estimates suggest it oegopsid squid. The aim of this study was to was 386 days old. Instantaneous growth rates better understand the biology of this species by were low (0.54% ML/day and 1.4% BM/day) but investigating its age and growth, as well as its consistent with enoploteuthid growth rates. When mating system. Toward this end, the age of 110 the growth rate of L. lorigera was corrected for individuals, ranging from 35 to 110 mm, was temperature encountered during the animal’s life, estimated by counting statolith growth incre- the growth rate was fast (0.47% BM/degree-days) ments. Estimates of age ranged from 131 to and consistent with the hypothesis that small 315 days and varied with mantle length. No cephalopods grow fast and that large cephalopods significant differences were found in the size of grow older, rather than fast. Mature females were often mated and had spermatangia in a seminal receptacle on the dorsal pouch behind the nuchal M. A. C. Roeleveld deceased. cartilage. Males probably transfer spermatangia to the females using their long second and/or H. J. T. Hoving (&) third arm pair since the paired terminal organs Department of Marine Biology, open far from the mantle opening. University of Groningen, CEES, P.O. Box 14, 9750 AA Haren, The Netherlands e-mail: [email protected] Keywords Lycoteuthis lorigera Á Growth rate Á Reproduction Á Ageing Á Statoliths M. R. Lipinski Á M. D. Durholtz Department of Environmental Affairs and Tourism, Private Bag X2, Roggebaai 8012, 8000 Cape Town, South Africa Introduction M. D. Durholtz e-mail: [email protected] Due to the full utilisation of neritic marine M. R. Lipinski resources, the exploitation of deep-water ecosys- e-mail: [email protected] tems is increasing. The collection of basic biolog- 123 260 Rev Fish Biol Fisheries (2007) 17:259–270 ical data on inhabitants of deep-water ecosystems The best method for estimating the age and is therefore important in designing management growth of squid is counting statolith increments. plans for sustainable exploitation of these systems. Daily deposition of increments has been validated One of the most important inhabitants of these for numerous species of squid, including oegops- ecosystems are squid because they form a trophic ids (for review see Arkhipkin 2004). Although the link between epipelagic and bathypelagic ecosys- daily deposition of increments has not yet been tems, as both active predators as well as important validated for L. lorigera, it has been in the prey items (Clarke 1996). Currently, little is known paralarvae of the enoploteuthid Abralia trigon- about the life history (age, growth and reproduc- ura, which belongs to a phylogenetic sistergroup tion) of most oceanic squid (Nesis 1995; Wood and of the Lycoteuthidae (Bigelow 1992; Young and O’Dor 2000; Arkhipkin 2004). Harman 1998). One particularly poorly understood oceanic The lifespan of oegopsid squid has been found family of squid is the Lycoteuthidae. This family to range from 3 months for the tropical Atlantic consists of two subfamilies and four genera that enoploteuthid Pterygioteuthis gemmata, which comprise a total of five described species. One reaches an adult size of 3 cm (Arkhipkin 1997), member of this family, Lycoteuthis lorigera to 2–3 years for the large sub-tropical mesope- (Steenstrup 1875), is an abundant squid in the lagic octopoteuthid, Taningia danae (Gonza´lez Benguela ecosystem. This species dominates, in et al. 2003). Most deepwater oegopsid squid, numbers the epibenthic cephalopod fauna on the however, tend to live for a minimum of 1 year continental slope of the South East Atlantic (Arkhipkin 2004). This is longer than the 1 year between 300 and 900 m depth (Roeleveld et al. or less lifespan of myopsids (reviewed by Jackson 1992). Although no current fisheries exist, the 2004). high abundance and muscular appearance of So far, no lycoteuthids have been aged. Given L. lorigera suggests that this species might be a its unique biology, it would be interesting to suitable resource for exploitation (Lipinski 1992). determine where L. lorigera will fit on the L. lorigera is preyed upon by commercially oegopsid longevity scale. It is generally accepted important fish, such as deep-water Cape hake, that squid that inhabit colder waters (either polar Merluccius paradoxus, and Kingklip, Genypterus or deeper waters) grow slower, and for longer capensis (Lipinski et al. 1992), as well as several (Arkhipkin 2004). To compensate for tempera- smaller cetacean species (Ross 1984), the Portu- ture differences, Wood and O’Dor (2000) intro- guese shark Centroscymnus coelolepis (Ebert duced the physiological timescale; in which et al. 1992), ribbonfish Lepidopus caudatus growth and longevity are corrected for temper- (Meyer and Smale 1991), the southern lantern- ature. Taking the animals’ mean life time tem- shark Etmopterus granulosus (Lipinski et al. perature of the environment into account, species 1992) and in New Zealand waters the species is that have a slow absolute growth rate can still preyed upon by petrels (Imber 1975). This species grow relatively fast. This approach will be used in feeds on pelagic crustaceans and fishes, including the interpretation of growth in L. lorigera. myctophids (Voss 1962). In addition, information will be presented on the In spite of the abundance and the important spermatophore production, morphology of the role of L. lorigera in the South East Atlantic male reproductive apparatus and the mode of ecosystem, little is known about the biology of spermatophore transfer from male to female, to this species. Clearly, this needs to be improved, better understand the mating system of the species. particularly as L. lorigera is a unique species amongst the oegopsids in having males that grow to a larger size than females. Another rare feature Materials and methods for squid is the presence of paired spermatophoric organs in males of L. lorigera. The aim of this An analysis of annual demersal surveys con- study is to investigate the age, growth and mating ducted by the Department of Environmental system of L. lorigera. Affairs and Tourism (DEAT) of the Republic of 123 Rev Fish Biol Fisheries (2007) 17:259–270 261 South Africa using the South African research images were recorded using AxioVision Ver. 4.2 vessel ‘‘Africana’’ and the Norwegian research software package supplied by Carl Zeiss Vision vessel ‘‘Dr. Fridtjof Nansen’’ between 1986 and GmbH. A number of over-lapping high resolution 2005 provided preliminary data on the distribu- images were taken of sectors with the highest ring tion of L. lorigera around South Africa. This clarity along transects from the nucleus, a distinct species was present in trawls between 192 and ring around the protostatolith, to the edge of the 1388 m deep, but was absent on the continental lateral dome (Fig. 3A). Increments were counted shelf (Fig. 1). from images using the counting feature of the L. lorigera used for ageing purpose were software. The rings in the outer part of the lateral collected during four research cruises in South dome (Fig. 3B) were often obscured, and extrap- African and Namibian waters (Table 1). The olation (based on the increment width of approx- mantle length (ML) and body mass (BM) of imately 10 of the last countable rings) was then animals were measured to the nearest mm and used to determine the number of rings in the 0.1 g respectively. The stage of sexual maturity periphery of the section. In order to increase was determined using the method of Lipinski and accuracy, counting the increments on every axis Underhill (1995). Statolith nomenclature was was repeated five times. The mean of the separate after Clarke (1978). Statoliths were extracted counts was used for that particular axis. and stored dry. The total statolith length (TSL) The instantaneous relative growth rate (G) was was measured to the nearest 0.01 mm. calculated using the equation: Statoliths (Fig. 2) were sectioned frontally G = (lnW2 –lnW1)/t, where W1 and W2 are through the lateral dome using the method of either ML (mm) or BM (g) at the beginning Lipinski and Durholtz (1994) and polished using and end of time interval t (Forsythe and Van polishing cloth. Sectioned statolith images were Heukelem 1987). recorded under phase contrast by an AxioCam Wood and O’Dor (2000) introduced the phys- MRc camera mounted on a Zeiss Axioscope 40 iological instantaneous relative growth rate (G/ compound microscope. A TV2/3 ‘‘C 0.63· camera T). G = (lnW2 –lnW1)/t2 – t1, where W1 is the mounted magnifying lens, together with a 10·/0.25 hatchling mass (g) and W2 is the mass of a mature or a 40·/0.65 objective lens, were used to produce female. t2 – t1 represents the age (days) at matu- low and high resolution images, respectively.
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