PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 119(1):91–136. 2006. of the Indian Ocean. A review. Part I. Inshore () collected during the International Indian Ocean Expedition

Patrizia Jereb and Clyde F. E. Roper* (PJ) Istituto Centrale per la Ricerca scientifica e tecnologica Applicata al Mare, Via di Casalotti 300, 00166 Rome, Italy, e-mail: [email protected] (CFER) Department of Invertebrate Zoology, National Museum Natural History, MRC 163, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A., e-mail: [email protected]

Abstract.—Cephalopods collected during the International Indian Ocean Expedition (IIOE; 1959–1965) and preserved at the Smithsonian Institution Oceanographic Sorting Center were analysed. This present work reports on the squids of the Loliginidae. A total of 378 specimens of Loliginids was identified: chinensis, Loligo duvaucelii, Loligo edulis, Loligo singha- lensis, Loligo (?) sumatrensis, hardwickei,andSepioteuthis lessoniana. A systematic description is provided for each , as well as its geographic distribution. Extensive updated information on the biology and fishery of each species also is discussed. Basic measurements were taken on each specimen and main morphological indices were computed; these were compared with those reported in the literature for each species. Tables provide measurements, morphological indices and comparisons with other studies. Appendices present information on capture sites, measurements and indices of individuals by sex and stage of maturity. These are located on the internet at www.biolsocwash/Jerebroper and http://www.nmnh.si.edu/iz/ cephs/morphology/

The International Indian Ocean Expe- agreement on the program was eventually dition (IIOE; 1959–1965) was a coopera- reached. It stressed the importance of tive, multidisciplinary, multiship scientific inter-calibration of data acquisition meth- venture (Wooster 1984), the need of ods and techniques. It was designed to which was first recognized during the obtain basic data on the geomorphology International Geophysical Year (IGY) in of the ocean bottom and to gather all 1957. The Scientific Committee on Ocean possible biological data and information, Research (SCOR) decided first to support from the baseline productivity of the a two-year international program of phys- waters to the main fisheries species as ical measurements in the waters of the deep well as the status of commercial fish oceans, then to investigate more compre- stocks for exploitation. hensively the deep waters of the Indian The Expedition witnessed the evolution Ocean during the third and fourth years. of ‘‘old time oceanography into new time In spite of the many different views on oceanography’’ (Aleem & Morcos 1984), the organization, objectives, and system- since oceanography experienced dramatic atic planning of the Expedition, an changes in techniques, instrumentation and ships during the six-year-span of the * Corresponding author. Expedition. Forty ships from 13 countries 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON participated, equipped with the most U.S.A. and 17 other countries (Wallen & advanced electronic devices. The initial Fehlmann 1974). The work of sorting and cost of the expedition was estimated at 12 dispensing specimens continues to the million dollars US; the final costs ex- present, although in a very much reduced ceeded 60 million dollars U.S. (Aleem & capacity in recent years. The SOSC Morcos 1984). function now is incorporated into the The fundamental observations made Smithsonian National Museum of Natu- in geology led to the elaboration of ral History, Department of Invertebrate the theory on seafloor spreading, which Zoology. then became a central unifying theory In spite of all the work done, 40 years in marine geophysics. The study of later, material still needs to be examined. the seasonally reversing currents associ- It is available for research by interested ated with the monsoon system stimulated scientists. the development of more elaborate The present work is the result of a joint research programs on the interaction research project between the Italian Con- between the oceans and the atmosphere. siglio Nazionale delle Ricerche (CNR) An example of such comprehensive pro- (National Research Council) and the grams is the Global Weather Experiment North Atlantic Treaty Organization carried out in 1979, which used satellite (NATO). This enabled the first author observations on a world-wide scale and to spend one year in Washington, work- pioneered modern satellite-based oceano- ing with the Smithsonian Institution and graphic studies. (Aleem & Morcos 1984). the Rosenstiel School of Marine and Among the results were important con- Atmospheric Sciences (Miami, Florida) sequences for local nations, such as to analyse some of the training programs for students and the material collected during the IIOE. origin of oceanographic studies. The Expedition also drew attention to the Materials and Methods high productivity of Indian Ocean waters, leading to an increased interest in the The bulk of material came from cruises development of sea fisheries in many of the research vessels R/V Anton Bruun countries; e.g., see Meiyappan & Mo- and R/V Ambariaka; a few additional hamed (2003). samples from other sources also were The phenomenal amount of biological examined. Once identified to species level, material gathered during the Expedition specimens were classified by an alphanu- required appropriate initiatives by par- merical code and information on sam- ticipating countries to process and dis- pling area, depth and gear were recorded. tribute it and to provide for appropriate Appendices 1A–G provide information research. In Washington, D.C., a working by species on locality of capture, sex and unit was established in 1962 as a mecha- number. nism to receive, sort and distribute to Measurements and indices (see Appen- specialists the mass of specimens expected dices 2A–G) used throughout this paper from United States participation in the are those given by Roper & Voss (1983), IIOE. This was the Smithsonian Ocean- using dorsal length (ML) as ographic Sorting Center (SOSC) (Wallen a standard. If the measure was repeated & Fehlmann 1974). About ten years after on symmetrical structures, ‘‘r’’ and ‘‘l’’ opening, SOSC had sorted nearly 3.7 indicate the right and left sides. The million specimens from over 4,500 sam- following additional parameters also were ples received, about 1.5 million of which recorded. Length had been sent to 140 specialists in the (NGL) was measured as an additional VOLUME 119, NUMBER 1 93 parameter to assess female maturity and (see Tables 1C to 7C for each species). Nidamental Gland Index (NGLI) was Specimens for which maturity stage or sex computed as the length of the gland as was not assessable were not considered in a percentage of mantle length. All mea- these tables. Table 8 summarizes mea- surements are in mm, if not otherwise surements and indices by sex for all seven indicated. species examined in this study. A three-stage maturity scale developed The published information available on for the , Illex coindetii (Jereb & the biology and fishery of each species is Ragonese 1995, see below), where 1 5 reviewed and summarized following the immature, 2 5 maturing, and 3 5 mature, systematic description and geographical was applied to describe sexual maturity in distribution sections. Nomenclatural ref- the specimens. No maturity stage was erences conform to those provided in assigned to specimens too small to be Sweeney and Vecchione (1998). sexed; these are referred to either as All Appendices and Tables are available ‘‘juveniles’’ or ‘‘unsexed’’ in the text. on the Internet at the following websites: Number and shape of teeth on sucker www.biolsocwash/Jerebroper and http:// rings of arm suckers and largest tentacu- www.nmnh.si.edu/iz/cephs/morphology/ lar club suckers also were recorded. Selected indices of biological parame- Results ters were chosen for each species accord- ing to sex, stage of maturity, and specific Class Cephalopoda Cuvier, 1797 locality in the Indian Ocean, to enable Subclass Bather, 1888 intraspecific comparisons when sufficient Superorder Decabrachia Boettger, 1952 material was available (see Tables 1 to 7, Order Teuthidea Naef, 1916 A and B). Also, our results were com- Suborder Myopsina d’Orbigny, 1841 in pared with analogous measurements tak- Ferussac and d’Orbigny, 1834–1848 en on the same species by other authors Family Loliginidae Lesueur, 1821

Macroscopic scale used to assess stage of maturity in the present material (from Jereb & Ragonese 1995:374) as presented in Table 3 and Appendix 2 (see on-line websites).*

Females Males Stage 1 NG are visible but thin and almost The gonad is small and inconspicuous. The (Immature) transparent. The ovary is filamentous testis and the SO are hardly visible. The and transparent. penis is a small appendage protruding close to the left gill. not formed. Stage 2 NG are whitish and well formed but do The testis is a white, flat, lanceolate body. (Maturing) not cover the underlying visceral mass. The SS is well-developed and may contain The ovary is conspicuously developed a few spermatophores. The penis is clearly and granular but does not occupy all visible. Hectototylus almost completely the posterior half of the mantle cavity. formed. The ova are visible. Stage 3 NG large and ripe, milk-white, almost The testis is conspicuous, fusiform and (Mature) completely cover the visceral mass. slightly yellowish. The SS is filled with Ovary very large, occupies fully half of spermatophores. A slight pressure is the mantle cavity. Anterior portion of sufficient to force them to extrude from the ovary and the oviducts filled with the penis. eggs that are amber-coloured in fresh specimens.

*NG5 Nidamental gland; SO 5 Spermatophoric organ; SS 5 Spermatophoric sac. 94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Distinctive features.—Transparent skin (Loliolus), Loliolus (Nipponololigo), Lo- (corneal membrane) covers eye lens. Fun- ligo (Loligo)andLoligo (). nel-locking apparatus a simple, straight However, in the years following the groove and ridge. Fins attach to lateral 1988 Workshop, new observations and regions of mantle. Arm suckers in two cladistic analyses on morphological (e.g., longitudinal series. Tentacular clubs with Alekseev 1989, Anderson 1996) and suckers in four series (except two series molecular (e.g., Brierley et al. 1996, in manal region of Anderson 2000) characters added new clubs). Hooks never present. Buccal con- parameters to the still-problematic status nectives attach to ventral margins of within the family. Another workshop on ventral arms, arms IV. Seven buccal loliginid systematics was convened during lappets possess small to minute suckers the CIAC 2003 meeting in Thailand. One (except in Pickfordiateuthis and Sepio- goal of the workshop was to resolve teuthis). Left ventral arm usually hecto- differences in generic-level classification cotylized in males; structure of the of the family. The loliginid taxonomists modified portion of the hectocotylus use- reached consensus that a new classifica- ful as a diagnostic character in most tion is appropriate, which includes ten species. Eggs spawned in gelatinous fin- genera and nine subgenera as valid ger-like egg masses attached to substrate. (Vecchione et al. 2005). Additional sub- Color usually reddish-brown, darker dor- genera remain undescribed or undeter- sally, but quite variable depending on mined. The revised classification is: species and behavioural situation. Loligo, Afrololigo, Alloteuthis, Dory- Remarks.—The systematics of the fam- teuthis (), Doryteuthis (Amer- ily Loliginidae [seven genera, 41–50 spe- igo), , Loliolus (Loliolus), cies, Nesis (1987)] recently has been the Loliolus (Nipponololigo), object of studies to clarify the position of (Lolliguncula), Lolliguncula (Loliopsis), some of its species and genera (Cohen Pickfordiateuthis, , 1976; Natsukari 1976, 1983, 1984a, (Uroteuthis), Uroteuthis (Photololigo), 1984b; Lu et al. 1985; Natsukari & Uroteuthis (Aestuariolus). Okutani 1975; Brakoniecki 1986; Alek- Since these recent nomenclatural seev 1989, 1991; Korzun & Alekseev changes are rather complex and not yet 1991; Vecchione et al. 1998; Vecchione universally accepted, we choose to use the et al. 2005). former, long-standing classification sum- A noteworthy testimony to the impor- marized by Nesis (1987), as these generic tance of the many unsolved systematic and specific names are well established problems is that one of the main sections both in the scientific literature and in the of the 1988 Cephalopod International fisheries. The newly proposed classifica- Advisory Council (CIAC) Workshop tions of Vecchione et al. (2005), however, (Washington, July, 1988) was devoted are reported within brackets as a matter entirely to the Family Loliginidae. The of information. decisions made during the Workshop Loligo Lamarck, 1798 required some modifications to the tradi- Subgenus Loligo Lamarck, 1798 (s.s.) tional classification (Vecchione et al. Loligo chinensis Gray, 1849 1998) that tentatively assigned the nine [Uroteuthis (Photololigo) chinensis,Vec- recognized groups of species into five chione et al. 1998, Vecchione et al., 2005] genera and four subgenera. These are Fig. 1A–J Sepioteuthis, Lolliguncula (Lolliguncula), Lolliguncula (Loliopsis), Uroteuthis (Uro- Loligo chinensis Gray, 1849: 74 (original teuthis), Uroteuthis (Photololigo), Loliolus description); type locality: China.— VOLUME 119, NUMBER 1 95

20* 40* 80* mo* 100* 120* 140*

fm^k r) 40°

20*

-40*

Map 1. Geographical distribution of Loligo chinensis, (shaded areas); black dots, present material, site- specific localities.

Tryon, 1879: 145 (re-description of the Photololigo chinensis, Natsukari, 1984a: type).—Natsukari & Okutani, 1975: 85, 232.—Dunning, 1998: 774. text-Figs. 1–4 (re-description of the Loligo (Photololigo) chinensis, Okutani, type).—Okutani, 1980: 33, Fig. 38.— 1990: 61, Fig. 74. Nesis, 1982: 141, Figs. 37i–k; 1987: 151, Uroteuthis (Photololigo) chinensis,Vec- Figs. 37i–j.— Roper et al. 1984: 86.— chione et al. 1998: 218.—Vecchione et Brakoniecki, 1986: 28–29.—Okutani et al. 2005. al. 1987: 97. Loligo indica, Hoyle, 1886: 156, pl. 26 FAO vernacular names.—En-Mitre Fig. 1–10 (non Pfeffer 1884); type squid; Fr-Calmar mitre; Sp-Calamar mi- locality: Arafura Sea, south of Papua trado. New Guinea. Type locality.—Canton fish market, Loligo etheridgei Berry, 1918: 243, Figs. China. 28–38, pls. 67–69, 1 table, type locality: Distribution.—Loligo chinensis is an Australian Sea.—Adam, 1954: 136, Indo-Pacific species (Map 1). Its distribu- Figs. 7–9, 1 table, Dobo, Aru Is., tion extends from the western Pacific: Arafura Sea. Japan, South China Sea by Gray (1849), Loligo formosana Sasaki, 1929: 109, text- Wakiya & Ishikawa (1921); Hong Fig. 161, pl. 30, Fig. 13, 1 table, type Kong by Voss & Williamson (1971), locality: Tainan market.—Voss & Wil- as L. formosana; Philippines by Voss liamson, 1971: 58, Figs. 19, 20, 22, (1963), as L. singhalensis; Indonesia by pl. 14, Hong Kong. Adam (1954) as L. etheridgei; northern, Loligo singhalensis, Okutani, 1980.—Ne- western and eastern Australian waters sis, 1982, 1987. by Berry (1918), as L. etheridgei; Yeat- Doryteuthis singhalensis, Voss, 1963.— man & Benzie (1994); to the eastern Voss & Williamson, 1971.—Roper et Indian Ocean: Andaman Sea, Thailand al. 1984.—Silas, 1986. by Natheewathana (1992); Bay of 96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. A–G. Loligo chinensis Gray, 1849. A, ventral view (Roper et al. 1984). B, dorsal view (redrawn from Roper et al. 1984; as Loligo singhalensis). C, tentacular club (Roper et al. 1984). D, tentacular club sucker ring (Roper et al. 1984). E, tentacular club sucker ring (Roper et al. 1984; as L. singhalensis). F, arm III sucker ring (Roper et al. 1984). G, arm III sucker ring (Roper et al. 1984, as L. singhalensis). H–J. Loligo chinensis Gray, 1849. H, hectocotylus (Dunning and Lu 1998). I, hectocotylus (Roper et al. 1984, as L. singhalensis). J, (Nateewathana 1992).

Bengal in the present work; Chikuni Appendix 1A for detailed locality and (1983), Silas (1986) as Doryteuthis singha- capture data. All measurements and lensis. relative indices are reported in Appendix Material examined.—Forty-five speci- 2A. Table 1A gives data on measurements mens of Loligo chinensis (24 males, 21 and indices of morphometric characters females) from the eastern side of the Bay by sex and stage of maturity. Selected of Bengal (IIOE) were examined. See ranges of indices are reported in Table 8. VOLUME 119, NUMBER 1 97

Fig. 1.—Continued.

Diagnosis.—Fin length in adults greater present; Nesis (1987), as Loligo chinensis, than 60% of mantle length (Fig. 1A, B). absent; Okutani (1980), as L. chinensis, Hectocotylized portion of the left arm IV present; present material, absent. from 33% to 50% of total arm length The fins are rhombic, with anterior and (Fig. 1H, I). Arm sucker rings with 10–15 posterior margins almost straight (poste- stout, pointed, conical teeth distally, the rior margin only slightly concave). Fins are proximal margin smooth; occasionally longer than wide, become more accentu- with rudimentary teeth only (Fig. 1F, ated with growth; fin length reaches 0.67 of G). Largest club sucker rings with 20–30 mantle length, fin length index (FLI) up to teeth, very unequal in size: large teeth 65–70%; fin width is about half the length (about ten) alternate with small, some- of the mantle, fin width index (FWI) up to times very small, teeth (Fig. 1D, E). 58%, present work. Gladius rather narrow: gladius width The head is stout, slightly narrower 6–8% of gladius length (Fig. 1J). One than the mantle at the mantle opening. present on each side of the The arms are muscular, moderately long . (longest arm attains 35% of mantle length Description.—The mantle is cylindrical (Okutani et al. 1987); the arm formula is and tapers posteriorly to a bluntly conical 3.4.2.1 (Natsukari & Okutani 1975). tip; mantle width 20–30% of ML, widest The arm suckers are biserial and of in its anterior part. A cutaneous ridge may medium size. Proximal suckers on arms II be present or absent on the ventral surface and III in males slightly enlarged (Brako- of mantle in adult males: Voss & William- niecki 1986). Chitinous sucker rings bear son (1971), as Dorytheuthis singhalensis, from 10–15 stout, acutely pointed, conical 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON teeth on the distal margin. The proximal 1993a) reports a year-round spawning margin is smooth, but sometimes bears season with two main peaks (March–June rudimentary ‘‘teeth-like’’ projections. The and August–November), a fecundity of left arm IV is hectocotylized in males in 3000–11,000 eggs and a length at 50% its distal half by the modification of maturity of 160 mm and 140 mm mantle suckers and stalks into fleshy, cone- length (ML) for males and females, shaped papillae; hectocotylized arm index respectively. Subsequent age analysis re- (HcLI) up to 70%; mature males have vealed that L. chinensis matured earlier a secondary sexual modification of en- during the warmer summer period than larged proximal suckers on arms II and in the cooler winter period, suggesting III (Brakoniecki 1986). Suckers on the that maturity is governed more by in- ventral series are slightly larger than those dividual size than by age (Jackson 1993). on the dorsal series. Probably one of the largest loliginid The tentacles are moderately long and species of the Indo-Pacific region, its slender. Clubs are slender and rather long; mantle length has been reported up to club length index (ClLI) 20–35% (Fig. 1C). 490 mm for males and 310 mm for Club suckers quadriserial, those of the females (Voss & Williamson 1971, as D. median rows slightly larger (1.2–1.5 times singhalensis), and a length-weight rela- larger, Natsukari & Okutani 1975, Okutani tionship was computed for the population et al. 1987) than those of the lateral rows. from the Gulf of Thailand (Chotiyaputta The large suckers of the bear 20–30 1993b). conical, stout, sharp teeth around the Our material consists of L. chinensis whole margin, greatly variable in size; specimens caught during the month among these, about 10 large teeth alternate of March at depths between 36 and with smaller, and even very small, incon- 81 m; no specimen was sexually mature, spicuous teeth (0–4) in a line. although several were in a maturing Buccal lappets with small suckers. stage. A pair of is present on the Fishery.—A commercially important ink sac, one on each side. Based on this species throughout its range (Roper et character alone, the species should be al. 1984), L. chinensis is reported to be included in the genus Photololigo (Natsu- exploited currently in the Yellow Sea and kari, 1984a). East and South China Sea, and it The gladius is rather narrow; gladius width probably constitutes, together with L. 6–8% of gladius length (Okutani et al. 1987). edulis, the majority of the Chinese squid The relative indices obtained for this catch (Guo & Chen 2000). It is quite species in different studies, as well as data extensively exploited by the Hong Kong in the present work, are reported in fishery (Voss & Williamson 1971, Oku- Table 1C for comparison. tani 1980, Chikuni 1983) and the Taiwan- Biology.—The biology of L. chinensis is ese trawlers. still poorly known, and most basic bi- Of occasional commercial relevance in ological information is lacking (see also Japanese waters (Voss 1973), L. chinensis Natsukari & Tashiro 1991). A neritic is likely to represent a small part of the species, L. chinensis has been reported squid catch around the Philippine Islands, within a depth range of 15–170 m, and its as well, and it constitutes about half of spawning period is likely to extend the trawl catch of squid in the South throughout the year (Roper et al. 1984, China Sea (Chikuni 1983, Chan & Noor as D. singhalensis). 1986). A thorough review of the cephalopod Loligo chinensis is one of the most resources of Thailand (Chotiyaputta important species for the Gulf of Thai- VOLUME 119, NUMBER 1 99 land fishery (Chotiyaputta 1993a, Chan- Loligo duvaucelii d’Orbigny, 1835, in tawong & Suksawat 1997). It also is likely Ferussac and d’Orbigny 1834–1838 to constitute a substantial portion of the [Uroteuthis (Photololigo) duvaucelii,Vec- catch in the Malaysian waters (Chikuni chione et al. 1998, Vecchione et al. 2005] 1983), where it is reported as the domi- Fig. 2A–E nant squid species for some local fisheries (Ashirin & Ibrahim 1992). Probably it is Loligo duvaucelii d’Orbigny, 1835, in also well represented in the Indonesian Ferussac and d’Orbigny, 1834–1838: fisheries (Arafura Sea, Java Sea; Chikuni 318, pl. 14, pl. 20, Figs. 6–16.—Adam, 1983), and it is one of the most important 1954: 132–136, Figs. 5–6.—Voss & commercial cephalopod species in north- Williamson, 1971: 60–62, pl. 17, Figs. ern Australian waters (Dunning 1982), 19, 20, 22.—Okutani, 1980: 35, Fig. 42. along with L. singhalensis. ?Loligo sumatrensis, Brock, 1887: 595 (in As far as the Indian Ocean fishery is Adam 1954). concerned, this species is well represented in Loligo duvauceli d’Orbigny, 1835, 1839, squid catches from the southeastern waters 1848; Voss, 1963: 71–74, Fig. 12.— of the Bay of Bengal (Thai coasts, Anda- Adam, 1973: 18–19, Figs. 1–3; Nesis, man Sea; Chikuni 1983, Chotiyaputta 1982: 145, Fig. 37s–u.—Roper et al. 1993a, Chantawong 1994). It is reported 1984: 87–88.—Brakoniecki, 1986: 29– among the commercially exploited squid 30.—Jothinayagam, 1987: 43–46, Fig. species of India (Silas 1986, as D. singha- 15.—Nesis, 1987: 154, Fig. 37 q–s.— lensis) and Sri Lanka (Chikuni 1983). This Nateewathana, 1992: 7–10, Fig.2. species also is assumed to be present in Loligo indica Pfeffer, 1884: 64, Figs. 3, the Arabian Sea (Chikuni 1983, Siraimee- 3a.—Goodrich, 1896: 7, Figs. 20–28.— tan 1990, as D. singhalensis), even though Massy, 1916: 218, pl. 23, Fig. 9, pl. 24, it is not clear what proportion it con- Fig. 11. tributes to the fishery landings there Loligo galatheae Hoyle, 1885: 183–184, because of the confusion in the loliginid 1886: 159–160, pl. 27, Figs. 1–12. nomenclature. ?Loligo oshimai Sasaki, 1929: 123–125, Remarks.—After its original descrip- pl. 30, Fig. 14, text Figs. 162–164. tion, this species was re-described several Photololigo duvaucelii, Natsukari, 1984a: times under different names, as summa- 231.—Dunning, 1998: 775. rized by Natsukari & Okutani (1975), Loligo (Photololigo) duvaucelii, Okutani, who also give a re-description of the type- 1990: 61, Fig. 75. specimen. Those authors synonomized L. Uroteuthis (Photololigo) duvaucelii,Vec- indica Hoyle 1886, L. etheridgei Berry, chione et al. 1998. 1918, and L. formosana Sasaki, 1929 with L. chinensis. FAO vernacular names.—En - Indian Our specimens agree fairly well with the squid; Fr - Calmar indien; Sp - Calamar re-description provided by Natsukari & indico. Okutani (1975), even though many speci- Type locality.—India and other sites. mens are in poor condition. The long- (Indo-Pacific: Sumatra, coast of Malabar, term confusion inherent in the convoluted Bombay, Pondichery, Batavia, Moluccas, regional loliginid nomenclature makes it fide d’Orbigny 1835 and Adam 1973). very difficult to assess the true importance Distribution.—An Indo-West Pacific of this species to the fisheries. However, it species (Nesis 1982, 1987; Roper et al. seems probable that L. chinensis is under- 1984), Loligo duvaucelii extends its distri- reported, at least in the western areas of bution throughout the Indian Ocean its range. (Map 2), from the South African coasts 100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Map 2. Geographical distribution of Loligo duvuacelii, (shaded areas); black dots, present material, site- specific localities; open circles, present material, general localities.

(Angola, Mozambique by Adam 1962), the Java and Arafura Seas (Chikuni the Red Sea (Adam 1973) and the 1983). Arabian Sea (Sarvesan 1974, Perera Material examined.—208 specimens of 1975, Oommen 1976, Chikuni 1983, L. duvaucelii (65 males, 132 females, 11 Kasim 1985, Silas 1986, Rao 1988, juveniles) were examined from Indian Siraimeetan 1990, Vidyasagar & Des- Ocean, mostly IIOE: 24 from the Arabian mukh 1992, Mohamed 1993, 1996), east- Sea, eight from Djibuti (Somalia), 52 ward to the Bay of Bengal, Sri Lanka from Madagascar and 124 from the Bay (Voss 1973, Sarvesan 1974, Chikuni 1983, of Bengal. See Appendix 1B for detailed Silas 1986, Mohan and Rayudu 1986, localities and capture data. All measure- Jothinayagam 1987, Jasmine et al. 1989) ments and relative indices are reported in and the Andaman Sea (Voss 1973, Appendix 2B. Table 2A, B give general Chikuni 1983, Nateewathana 1992, Cho- and regional data on morphometric tiyaputta 1993a, 1997). A common Ma- measurements and indices by sex and laysian squid (Sumatra, Malaysia, Thai- stage of maturity. A comparison of land by Voss & Williamson 1971, Voss measurements and indices reported by 1973, Latif 1982, Chikuni 1983, Chotiya- various authors is provided in Table 2C. putta 1993a), L. duvaucelii also is very Selected ranges of indices are reported in abundant in Philippine waters (Voss 1963, Table 8. Voss & Williamson 1971, Chikuni 1983) Diagnosis.—Fin length in adults up to and moderately abundant in the South 60% of mantle length (see Table 2C) China Sea (Shin 1982, Chikuni 1983). (Fig. 2A). More than half the length of It has also been recorded in the West the left ventral arm hectocotylized in Pacific north to Formosa Island and males, up to 75% in present work (see the Taiwan Strait (Adam 1962, Voss Table 2A, C) (Fig. 2D). Suckers of lateral 1963, Roper et al. 1984) and south to arms of males (greatly) enlarged. Arm VOLUME 119, NUMBER 1 101

Fig. 2. Loligo duvaucelii d’Orbigny, 1835. A, dorsal view (Roper et al. 1984). B, tentacular club sucker ring (Roper et al. 1984). C, arm III sucker ring (Roper et al. 1984). D, hectocotylus (Nesis 1982/1987). E, gladius (Voss & Williamson 1972).

sucker rings with broad, large, square (photophores) present along the ink sac, teeth (five to nine) on the distal margin in one on each side. females and around the entire ring (up to Description.—When in good condition, 18) in males (Fig. 2C). Club sucker rings our material conforms to the general with 14 to 20–22 short, sharp teeth descriptions given by Adam (1954) regularly spaced around the entire margin and Voss (1963). A more delicate appear- (Fig. 2B). Two oval luminous organs ance (i.e., more slender mantle, smaller 102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON head), however, is evident in the speci- cotylized arm is unusually short. Two mens from Djibuti (Somalia) (Table rows of fleshy papillae occur distal to the 2B-2). suckers. They vary in size and form: those Mantle moderately long and slender, of the ventral row are cone-shaped, cylindrical for about half (slightly more sometimes very ‘‘fat’’ and long (e.g., than half) its length, then it tapers gently present material, Bay of Bengal). Gener- into a blunt tip. Anterior margin with ally they are fused with the protective a small rounded lobe in the dorsal mid- membrane for at least their basal portion line. (more generally up to 0.33 of their Fins gently rhombic, broad, widest length). The papillae of the dorsal row approximately at the middle point of usually are thinner and shorter. Minute their length; fin length occupies more suckers are present on the tips of these than half of mantle length (up to 60%). papillae. The modified portion extends The anterior and posterior margins are for more than half of the arm length, up nearly straight, the lateral angles very to 75% in some of our material rounded. (Tables 2A, C). A peculiar feature is Head broad, slightly narrower than the noted in some of the specimens we mantle; eyes large. examined: at the base of the arm the Arms moderately long; arm formula dorsal protective membrane is slightly 3.2.4.1 (Voss 1963). Sexual dimorphism is expanded to form a kind of crest that noticeable in sucker size and dentition, enfolds the first basal suckers. This crest even though not always as strongly is present in a few specimens from the evident in the material we examined as southwestern and southeastern coasts of stressed by other authors (e.g., Voss India, i.e., off Cochin (PJ 308 and 309) 1963). and off Mandapam (PJ 313). The suckers of all arms in females are Tentacles long; clubs large, up to 45– similar in size and dentition: 4–10, usually 50% of ML; club suckers quadri-serial, 6–7, broad, square, bluntly edged teeth those of the medial rows slightly larger occur along the distal margin whereas the (about 1.5 times) than those on the lateral proximal margin is smooth. Very often rows. Sucker rings have 14–22 acute, the central tooth (teeth) is (are) more conical, regularly spaced teeth around slender and slightly pointed, as evidenced the entire margin. Smaller, almost in- also in the present material. As a second- conspicuous denticles may be present ary sexual modification in males, suckers between adjacent conical teeth. of arms II and III usually are enlarged, Buccal membrane seven-pointed; each sometimes greatly so, and sucker rings lappet bears up to 5 small suckers. bear several [usually around ten, but up A pair of ovoid luminous organs to 18 according to Adam (1954)] broad, (photophores) is located on the ventral square, truncate teeth that occupy almost side of the ink sac, near the anus. Details the entire margin. These teeth are shorter of the morphology and physiology of than those of females. Small, inconspicu- these organs are presented by Prinngen- ous denticles may be present along the nies & Jorgensen (1994) and Prinngennies proximal margin (Adam 1973). et al. (2001). On the basis of this Left ventral arm hectocotylized in character, the species should be included males. Usually about half of the arm in the subgenus Photololigo Natsukari bears from 12–22 pairs of normal suckers; (1984a). up to 30 pairs are present on our speci- Gladius with a long, straight edged mens from the east African coast, in vane in mid-portion and a short rachis which the modified portion of the hecto- (Fig. 2E). VOLUME 119, NUMBER 1 103

Biology.—In recent years, increased studies have confirmed an allometric attention has been paid to the biology of growth (Silas 1986, Mohamed 1996, L. duvaucelii, principally because of the Karnik & Chakraborty 2001), with fe- importance of the fishery for this species males growing faster than males; howev- in the Indian (both west and east coasts) er, males ultimately attain a greater size and Thai waters. A list of local names of and age. Maximum reported size of males this species in Indian waters is given in is 320 mm ML from Thai waters (Cho- Meiyappan & Mohamed (2003). tiyaputta 1993a) and 330 mm ML along While its embryonic development and the west coast of India (Meiyappan & hatching were observed in the laboratory Mohamed (2003). (Asokan & Kakati 1991), the biology of Recently, a first attempt to culture the species by field data and information L. duvaucelii was made from eggs was studied from the Indian waters by collected in the field (Prinngennies et al. Oommen (1976), Silas (1986), Rao (1988), 2000). Hatchlings were fed with rotifers Vidyasagar & Deshmukh (1992), Mo- and Artemia and survived for several hamed (1993, 1996), Mohamed & Rao weeks. (1997), Rahim & Chandran (1984), Kar- Fishery.—Loligo duvaucelii is one of the nik & Chakraborty (2001), Karnik et al. most common species among the Indo- (2003). Field studies also were conducted Pacific loliginids, and it was reported to from the Gulf of Thailand and the be the most important squid species for Andaman Sea by Chotiyaputta (1982, the Indian fishery during the years 1972– 1993a, 1993b, 1994, 1997), Supongpan 1984 (Ramachandran 1987). It was con- (1984, 1988), Manoch & Shunji (1992) sidered the most promising species for the and Supongpan & Sinoda (1998). Indian dried squid industry (Sarvaiya According to information from stato- 1991), and it constituted, along with L. lith age readings (Chotiyaputta 1997), chinensis, about 90% of the squid catches the life cycle of L. duvaucelii would be of Thailand during the year 1988, prob- about one year, which, as expected, is ably a typical situation for that area shorter than estimates obtained by length (Chotiyaputta 1993a). frequency analysis (e.g., Kasim 1985, Known to be fairly important for the Mohamed 1996, Mohamed & Rao fisheries of the eastern Arabian Sea 1997). The spawning period appears to (Chikuni 1983), this species constituted be quite prolonged, almost all year round, 68% (i.e., 4795 metric tons) of the squid with peaks in different months, principal- caught north of and off the Gujarat coast ly in spring and autumn. Spawning during December 1983–March 1984 aggregations occur in the post-monsoon (northwest coastal India, Siraimeetan months along the west coasts of India 1990). The catch continued at a median (Mohamed 1993) and seem to be de- annual value of over 5000 metric tons pendent upon the SW and NE monsoons into the middle 1990’s (Mohamed & Rao in the western part of the Gulf of Thai- 1997). It formed about 8% of the trawl land (Supongpan & Sinoda 1998). Size at landings at Bombay with a standing stock 50% maturity is 90–130 mm ML for of 990 metric tons and an annual stock of females and 70–150 mm ML for males. 2150 metric tons, according to Vidyasa- Observations on growth after sexual gar & Deshmukh (1992). In this area, maturity is reached indicate an extended (Sepiidae) constitute the domi- reproductive phase within the life cycle, nant species among cephalopods for all i.e., a non-strictly semelparous reproduc- capture techniques (Siraimeetan 1990), tion (Mohamed 1993), as is the case in but L. duvaucelii represented 68% of the other squid (Rocha et al. 2001). Other catches of the Mangalore trawlers of 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON central west India (Silas 1986). The most common squid species caught in the species is present in small quantities Java Sea (Sudjoko 1987). throughout the year in the trawl fishery Loligo duvaucelii is abundant in the off Cochin, southwest India, and it is the South China Sea (Chikuni 1983, Guo & dominant squid caught as a by-catch by Chen 2000), and it is reported among the the shrimp trawlers (Silas 1986). commercial squid species of the Hong Loligo duvaucelii also is reported to Kong fishery (Shin 1982). However, very constitute a significant fraction of the little information is known about the total cephalopod production of Vizhin- abundance and landings of this species jam, southwestern India (i.e., about 42%, in the Yellow and East China Seas Silas 1986), where it is caught by shore (Chikuni 1983). seine, boat seine, and hook and line. Loliginid squids constitute 48% of the Loligo edulis Hoyle, 1885 total Indian cephalopod captures versus [Uroteuthis (Photololigo) edulis,Vec- 51% for sepiioids (1% for octopods). chione et al. 1998, Vecchione et al. 2005]. Loligo duvaucelii is by far the predomi- Fig. 3A–H nant squid species in that total, represent- ing 49% of the west coast cephalopod Loligo edulis Hoyle, 1885: 186; 1886: 152, landing (20–330 mm ML) and 24% of pl. XXIII, Figs. 1–9.— Sasaki, 1929: 107– east coast landings (30–180 mm ML). 109, Figs. 57–59, pl. XIII, as form nagasa- Peak Indian cephalopod landings oc- kiensis and form grandipes.—Voss, 1963: curred in 1995 at 116,753 metric tons 67, Figs. 11a–e.—Voss & Williamson, and averaged around 110,000 metric tons 1971: 57, Figs. 19,20,22, pls. 14–15.— from 1996 through 2000 (Meiyappan & Adam, 1973: 19, Figs. 4–5.—Okutani, Mohamed 2003). 1980: 33, Figs. 36–37.—Nesis, 1982: 141, As for the Bay of Bengal, L. duvaucelii Figs. 37a–e, 1987: 151, Figs. 37b–f.— is important for all the main eastern Roper et al., 1984: 88.—Brakoniecki, Indian landing places (i.e., Waltair, Ka- 1986: 27–28.—Okutani et al., 1987: 99. kinada, Madras, Mandapam, Vishakha- Loligo chinensis, Sasaki, 1914: 601.— patnam), where it constitutes the most Wakiya and Ishikawa, 1921: 279 (non abundantly caught squid species (Mohan Gray 1849). & Rayudu 1986, Silas 1986, Kripa et al. Loligo kensaki Wakiya and Ishikawa, 1996). It also is reported as the most 1921: 283, pl. 2, Fig. 9. common species landed along the Anda- Loligo budo Wakiya and Ishikawa, 1921: man Sea coast of Thailand (Natee- 258, pl. 2, Figs. 1,10.—Okada, 1927: wathana 1992, Chantawong 1994). 174, Fig. 2.—Roper et al. 1984: 89– In the Gulf of Thailand, L. duvaucelii is 90.—Nesis, 1982: 144, Figs. 37z, 37zh, caught by various fishing gears, such as 1987: 151, Figs. 37G–H.—Okutani et otter trawl, pair trawl, squid night-light al. 1987: 101. attraction, push net, and hook and line; in Loligo singhalensis, Adam, 1954 (fide 1988 the species contributed about 45% Korzun & Alekseev, 1991), non Ort- of the 67,176 metric tons of landed squid mann, 1891. (Chotiyaputta 1993a). Doryteuthis kensaki Wakiya and Ishi- Loligo duvuacelii is one of the main kawa, 1921: 283, Fig. 4, pl. 1: Fig. commercial squid species for the Philip- 9.—Okutani, 1973: 94, Fig. 40. pine fishery (Voss 1963, 1973; Chikuni Photololigo edulis, Natsukari, 1984a: 1983), and it is reported among the five 230.—Dunning, 1998: 776. major squid species known in the Malay- Loligo (Photololigo) edulis, Okutani 1990: sian area (Latif 1982). It is among the 57, Fig. 67. VOLUME 119, NUMBER 1 105

20" 40* w 00" 100* 120* 140*

fmJk 40°

••V c^^ 20° \; \*

* k -20"

-40°

Map 3. Geographical distribution of Loligo edulis, (shaded areas); black dots, present material, site- specific localities.

Uroteuthis (Photololigo) chinensis,Vec- Sea, Thailand, (Nateewathana 1992, Cho- chione et al. 1998: 318. tiyaputta 1993a) and the Bay of Bengal (Chikuni 1983, present work), to the FAO vernacular names.—En - Swordtip Arabian Sea, including the Gulf of Aden, squid; Fr - Calmar epee; Sp - Calamar the Gulf of Oman, and the Persian Gulf espada. (Zuev 1971, Voss 1973, Shvetsova 1974, Type locality.—Yokohama Fish Mar- Chikuni 1983, present work), the Red Sea ket, Japan. (Adam 1973), and southward to Mozam- Distribution.—An Indo-Pacific species bique (East African coast by the present (Nesis 1987), Loligo edulis is fairly com- study; Saja de Malha Bank by Zuev 1971, mon and relatively abundant in the Voss 1973; Mozambique Channel by western Pacific (Map 3), from northern Korzun 1992). waters (East China Sea, Japan Sea by Material examined.—Thirty-six speci- Okutani 1973, 1975, 1980; Okutani et al. mens of L. edulis (11 males, 15 females, 1987; Chikuni 1983; Natsukari & Tashiro 10 juveniles), 13 from the Bay of Bengal, 6 1991; Kubota et al. 1993; Zheng 1994) to from the Arabian Sea and 17 from the the tropical seas (Indonesia, Java Sea, coasts of East Africa (Indian Ocean, Malaysia, Thailand by Latif 1982, Shin IIOE). See Appendix 1C for detailed 1982, Chikuni 1983, Chan & Noor 1986, locality and capture data. All measure- Okutani et al. 1987, Sudjoko 1987, ments and relative indices are reported in Sodikin 1992, Chotiyaputta 1993a), south Appendix 2C. Data on measurements and to northern Australia (Okutani 1980, indices of morphometric characters by Okutani et al. 1987, Dunning 1982, sex and stage of maturity are given for Chikuni 1983, Yeatman & Benzie 1994). the total collection of study material The distribution of L. edulis also extends (Table 3A) and for regional areas throughout the Indian Ocean, from its (Tables 3B, 1–3). Selected ranges of indices southeastern waters, i.e., the Andaman are reported in Table 8. 106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Loligo edulis Hoyle, 1885. A, ventral view (Roper et al. 1984; as L. edulis form edulis). B, dorsal view (Roper et al. 1984; as L. edulis form budo). C, photophores (L.O.) on ink sac (Roper et al. 1984). D, tentacular club (Roper et al. 1984). E, tentacular club sucker ring (Roper et al. 1984). F, arm sucker ring (Roper et al. 1984). G, hectocotylus (Roper et al. 1984). H, gladius (Voss & Williamson 1972).

Diagnosis.—Fin length in adults ranges square-cut (bluntly-pointed) teeth on the from a little more than 50% up to 70% of distal margin; the proximal margin mantle length (see Table 3A) (Fig. 3A, B). smooth, or only irregularly denticulate Hectocotylized portion of the left arm IV with inconspicuous denticles (Fig. 3F). very long, up to 65–80% of its length Largest club sucker rings with up to 30– (Fig. 3G). Arm sucker rings with up to 40 teeth very unequal in size: usually a dozen (more often 6–8) long, slender, about 15–20 large, stout, sharp, conical, VOLUME 119, NUMBER 1 107 widely-spaced teeth, alternating with on ‘‘medium size specimens’’, Adam smaller ones, sometimes minute (from 1954) and even ‘‘cubic’’ (Okutani et al. one to groups of three to four) (Fig. 3E). 1987), with prominent eyes. Head width The gladius of ‘‘…usual form, the narrow varies between 10–11% and 20–37% of anterior portion being less than 0.25 of mantle length (Voss 1963, Okutani et al. the total length…’’ (Hoyle 1885: 187), has 1987). a broad rachis, with a large, oval vane Arms moderately long, 25–45% of with rounded margins (Voss 1963) mantle length. The arm formula is vari- (Fig. 3H). One luminous organ present able, 3.4.2.1 (Hoyle 1885, Sasaki 1929, on each side of the ink sac (Fig. 3C). Voss 1963, Okutani et al. 1987) or 4.3.2.1 Description.—As far as many charac- (Okutani et al. 1987). The suckers are ters are concerned, the following descrip- biserial, oblique and of medium-size, tion deals with very broad distributional sometimes slightly larger in males (Voss limits, due to the polymorphic character- 1963) and in the lateral arms (Adam istics of L. edulis (see Remarks and 1973). The suckers on arms II and III of Table 3A, B). mature males are noticeably enlarged as The mantle is moderately stout to a secondary sexual character (Brako- elongate, cylindrical anteriorly, gradually niecki 1986). Arm sucker rings bear up tapered posteriorly to a blunt point. to a dozen (more often 6–8) long, slender, Mantle width 30–40% of mantle length. mostly squared, sometimes conical, but Mature males may be more slender always flat-topped (blunt) teeth on the (Okutani et al. 1987) and often are distal margin. They can be widely spaced reported to bear a cutaneous ridge on or closely packed (also present material). their ventral mantle surface (Sasaki 1929: The proximal margin is either smooth or ‘‘…ordinarily present …’’; Adam 1954 (as slightly irregularly denticulate, with very L. singhalensis, alias L. edulis following low, rounded denticles (Adam 1973). Korzun & Alekseev, 1991, Roper et al. The left arm IV is hectocotylized in 1984, Okutani et al. 1987, Nesis 1987). males for more than 0.75 of its length This cutaneous ridge, however, is not (HcLI up to 70–80 in mature males, mentioned by Voss (1963) nor Adam Table 3A), by the modification of the (1973) in their description of specimens suckers and stalks into fleshy, conical from the Philippines and from the Red papillae, which may or may not bear Sea, respectively. No mature male was small suckers on the tips. On the proximal available in the present work, and no part of the arm from six to then up to 27– ridge was detected on the specimens 30 pairs of normal suckers are present. examined. The existence of different ‘‘forms’’ within Fins large, rhombic with the anterior the species L. edulis is based primarily on margin slightly convex, the posterior these variations. margin gently concave, and the lateral Tentacles moderately long, slender. angles rounded. Fins become slightly Clubs expanded, lanceolate; club length longer than wide in adult specimens (up varies between 15% and 40% of mantle to 70% of mantle length), their width length (Fig. 3D). Club suckers quadri- reaching 60% of mantle length (usually serial; medial suckers larger than the slightly larger in females). lateral ones (about 1.2 times greater The head seems quite variable in its diameter, Okutani et al. 1987). The largest relative dimensions, having been de- club sucker rings bear up to 30–40 teeth scribed alternatively as ‘‘large’’ (Sasaki around the entire margin, very unequal in 1929, on ‘‘full length specimens’’; Voss size: often 15–20 large, sharp, widely- 1963), ‘‘small’’ (Hoyle 1885, Sasaki 1929, spaced, conical teeth alternate with much 108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON smaller ones, sometimes rather regularly northwestern Pacific indicate a highly (one to four smaller between two larger, variable growth depending on season Okutani et al. 1987). and area (Natsukari & Tashiro 1991). Buccal lappets with several small suck- Data from northern Australia support an ers (from 2–3 up to14–15; Voss 1963, extended spawning season for the species Sasaki 1929, Adam 1954, 1973). A pair of and show that L. edulis in the Gulf of oblong photophores is present on the ink Carpentaria reaches sexual maturity at sac, one on each side. smaller sizes (70–80 mm ML) than the The gladius is long and moderately northern form (Dunning et al. 1994). narrow, 6–7 times longer than wide; the Loligo edulis in the northwestern part of vanes are distinctly curved laterally; the the Indian Ocean also is reported to reach rachis is about 0.2 the length of the sexual maturity at the small size of 70– gladius. 80 mm (Shvetsova 1974) and even smaller The relative indices obtained for L. edulis (50–60 mm, Adam 1973), and its - by different authors, as well as in the ing period extends from fall to spring or present work, are reported in Table 3C. even to the beginning of summer. Biology.—An extensive review of the Fishery.—A medium to large-sized status of the biology and resources of L. species, L. edulis has excellent flesh edulis in the Japanese waters was pub- quality, and it is exploited throughout lished by Natsukari & Tashiro (1991). its distributional range (Roper et al. According to those authors, this species 1984). One of the principal commercial inhabits continental shelf waters in Japan squid species of the Japanese market and winters close inshore in shallow (Wray 1996), it constitutes one of the water. The spawning season extends most important resources for the throughout the year, with three detectable coastal fisheries of Kiushu and the peaks in spring, summer and autumn, but southwestern Japan Sea, where the annu- the season is shorter in the northernmost al catch is estimated around 25,000 metric part of the area. The life span is reported tons. It is caught throughout the year by to be about one year (see also Natsukari jigging, set netting, bottom trawling and et al. 1988). Juveniles feed preferentially other gears (Natsukari 1991, Kubota et al. on crustaceans, whereas the main food 1993). for the adults is fishes. Loligo edulis also is very abundant in The size at which individuals become the Yellow and East China seas (Chikuni sexually mature shows great variability, 1983, Zheng et al. 1999), where, together depending on the season and locality. with L. chinensis, it is believed to account Most specimens of both sexes reach full for the majority of the Chinese squid maturity by 150–200 mm ML. The small- catch (Chikuni 1983, Zheng 1994, Song et est size recorded for full maturity was al. 1999, Ling & Zheng 2000). 52 mm and 59 mm ML for males and It is vigorously exploited by the Hong females, respectively, while some speci- Kong fishery, where it constitutes, to- mens were not sexually mature at a gether with L. chinensis, a large pro- size larger than 300 mm ML. Spawning portion (50–60%) of the annual cephalo- grounds were recorded at different depths, pod landings (Voss & Williamson 1971, from 30–40 m down to 100 m, on sandy Shin 1982), and it is believed also to be bottom in coastal waters where warm rather abundant in the South China Sea oceanic currents inflow. (Chikuni 1983). Loligo edulis is fished in The analysis of several length-weight the Malaysian waters (Latif 1982, Chan & relationships obtained for L. edulis in Noor 1986), and it represents one of the different seasons and locations of the main species of the Philippine and Indo- VOLUME 119, NUMBER 1 109 nesian fisheries (Voss 1973, Sodikin 1992, kensaki Wakiya & Ishikawa, 1921, and L. Sudjoko 1987). It also constitutes one of budo Wakiya & Ishikawa, 1921. the most important commercial cephalo- Three ‘‘forms’’ subsequently were iden- pod species in the northern Australian tified by Sasaki (1929): L. edulis (non waters (Dunning 1982). Hoyle, but corresponding to L. kensaki Loligo edulis is highly abundant in the Wakiya & Ishikawa, 1921); L. edulis form Andaman Sea, where it represents one of nagasakiensis (corresponding to L. edulis the main Thai squid resources (Chotiya- Hoyle, 1885), to which Sasaki referred putta 1993a), but it is not mentioned three specimens previously described by within the cephalopod resources of India him (Sasaki 1914) as L. chinensis Gray, (Silas 1968, Rao 1973, Perera 1975, Silas 1849; L. edulis form grandipes (corre- et al. 1982, Silas 1986, Mohan & Rayudu sponding to L. budo Wakiya & Ishikawa, 1986, Jothinayagam 1987, Siraimeetan 1921). Sasaki did not consider L. chinensis 1990). However, its presence in the (non Gray 1849) a valid form. western Indian Ocean (i.e., Arabian Sea, Okutani (1975), in the revision of his Red Sea, East African waters) was catalogue of Japanese squid, synony- confirmed (Zuev 1971, Adam 1973, Voss mized two of the three forms, erecting 1973, Shvetsova 1974, Chikuni 1983, two subspecies: L. edulis edulis Hoyle, Korzun 1992), even though its potential 1885 (corresponding to L. kensaki Wa- to the fishery there is still unknown. We kiya & Ishikawa, 1921 and L. edulis form suggest that the apparent absence of its nagasakiensis Sasaki, 1929) and L. edulis occurrence in Indian waters is a matter of budo Wakiya & Ishikawa, 1921 (corre- misidentification rather than of disjunct sponding to L. edulis form grandipes distribution. Sasaki, 1929). Remarks.—Loligo edulis is character- Natsukari (1984a:230), in one of his ized by a marked polymorphism, both by studies of loliginid squids, proposed the locality and by season (Okutani et al. new genus Photololigo, distinguished 1987, Nesis 1987, Natsukari & Tashiro from the other loliginids essentially by 1991). The existence of such a variety of the presence of luminous organs (photo- ‘‘forms’’ whose taxonomic relationships phores) on the ink sac, and he designated are still not clear, makes the confident L. edulis Hoyle, 1885 as the type species identification of the species rather com- of the new genus. Later Natsukari et al. plicated and the assessment of popula- (1986, 1988) stated the opinion that the tions (stocks) rather difficult. After the two subspecies recognized by Okutani original description by Hoyle (1885), who (1975) could be different seasonal referred to one male specimen from the ‘‘forms’’ of the same species, probably Yokohama Fish Market, Japan, the not genetically isolated, and that the condition of maturity of which was not differences between the two forms were specified, the species was repeatedly re- evident only in late (subadult) stages. described by subsequent authors, as Okutani et al. (1987:101) took into reported below and summarized in account these considerations but still Table 3D. referred to the two forms as valid, L. Among the so-called L. edulis-type edulis Hoyle, 1885 and L. edulis form squid distributed throughout the western budo Wakiya & Ishikawa, 1921. Okutani Pacific (Natsukari & Okutani 1975, Nat- et al. (1987:99) also clearly pointed out sukari & Tashiro 1991), four species that L. edulis Hoyle, 1885, is a species originally were recognized by Wakiya characterized by ‘‘…polymorphism by and Ishikawa (1921): L. edulis Hoyle, locality and by season within the range 1885, L. chinensis (non Gray, 1849), L. of the distribution….’’ 110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Nesis (1987) reported two species: L. Unfortunately, among those we exam- edulis Hoyle, 1885 distributed from the ined, only the specimens from the Arabi- Red Sea and Mozambique into the an Sea (2 males and 4 females, 48–65 mm western Pacific, and L. budo Wakiya & ML) were in good condition, but none Ishikawa, 1921 from Southern Japan. was fully mature. Their general morphol- However, he actually considered L. budo ogy closely resembles that of L. duvauce- to be a ‘‘doubtful species,’’ possibly lii, as already noted by Adam (1973) for a ‘‘form’’ of L. edulis (Nesis 1987: 151). his specimens from the Red Sea. From Also, referring to the northwestern Pacific these, L. edulis is distinguishable essen- population, Nesis (1987: 151) discussed tially by the arm sucker dentition: it two different forms ‘‘…of unclear taxo- strictly conforms to the L. edulis-type, nomic relationship…’’: L. edulis form with six to eight long, slender, blunt teeth edulis Hoyle, 1885 (includes L. edulis on the distal margin of the ring and by the form nagasakiensis Sasaki, 1929), and longer modified portion of the hectocot- L. edulis form kensaki Wakiya and ylized arm. The hectocotylized arm in our Ishikawa, 1921 (includes L. edulis form two males is very well developed in spite of edulis Sasaki, 1929, non L. edulis Hoyle, the small size of the specimens (48–65 mm 1885). ML, Table 3B-1; see also Adam, 1973). It While L. edulis form budo is character- bears six and eight pairs, respectively, of ized by very large clubs, long arms and normal suckers at the proximal base of the large suckers in comparison with the arms, followed by the sucker stalks mod- typical L. edulis, the two L. edulis ified into cone-shaped papillae with min- ‘‘forms’’ differ essentially in the hectocot- ute suckers on the tip, exactly as described ylized structure: 7–8 pairs of normal by Voss (1963). The club sucker dentition suckers at the base of the arm, no small conforms to the alternating large and small suckers on the papillae of the modified tooth pattern and, according to Adam portion (5L. edulis form edulis), versus (1973), ‘‘…hardly more than one small 20–27 pairs of normal suckers and tooth…’’ is distinguishable between two rudimentary suckers on the papillae of large teeth. the modified portion (5L. edulis form Our material from the East African kensaki). coasts (6 males, 5 females, 6 juveniles of Natsukari & Tashiro (1991) again undetermined sex, ML 30–87 mm) is in stated that they consider L. edulis form poor condition, but it also is morpholog- kensaki Wakiya & Ishikawa, 1921 and L. ically very similar to the appearance of L. edulis form budo Wakiya & Ishikawa, duvaucelii. The arm sucker dentition, in 1921 as two seasonal forms of the same some cases (i.e., specimens 154 n.1–2 and species, based on some evidence that the 159 n.1), is very close to the chinensis- forms probably are not genetically iso- type, in the sense that the teeth are rather lated (Natsukari et al. 1986, 1988). conical and appear sharply pointed; un- Subsequently, four species of Photolo- der higher magnification, the teeth have ligo from the northern waters of Australia blunt tips. Other specimens have an were identified using allozymes (Yeatman obvious edulis-type dentition, with long, & Benzie 1994). Two of these species squared, widely spaced teeth. Also, the fitted the gross morphology of Photolo- hectocotylus is rather variable, with ligo edulis from Japan, supporting the regard to the number of normal suckers opinion that a number of widely distrib- at the base of the arm, between 11–12 and uted species (such as P. edulis)may 16–18 pairs. The morphology of the cone- represent a series of morphologically shaped papillae, whether longer and more similar allopatric sibling species. slender or shorter and more stout, always VOLUME 119, NUMBER 1 111 includes small suckers on the tips. The Loligo sumatrensis d’Orbigny, 1835, in club sucker rings have 10–11 large teeth Ferussac and d’Orbigny 1834–1838 that alternate with 1 or more (two to [Loliolus (Nipponololigo) sumatrensis, three) small, inconspicuous teeth. Vecchione et al. 1998; Vecchione et al., The specimens from the Bay of Bengal pers. comm.]. (3 males, 6 females, 4 juveniles of un- Fig. 4A–I determined sex) also are in poor condi- tion. They appear to be morphologically Loligo sumatrensis d’Orbigny, 1835, in quite different at first glimpse, almost Ferussac and d’Orbigny 1834–1838: indistinguishable from our L. chinensis 317–318, Loligo pl. 13, Fig. 1–3.— specimens from the same area, with Natsukari, 1984b: 259–263, Figs. 1–2, slender mantles, smaller heads, and slight- tab.1.—Okutani et al. 1987: 91.—Na- ly more narrow and more ‘‘rhombic’’ fins teewathana, 1992: 14–17, Fig. 4, tab. 5. (clearly longer than wide). It is impossible Loligo kobiensis Hoyle, 1885: 184–186.— to know how much the variations are Roper et al. 1984: 94.—Brakoniecki, ascribable to effects of fixation. This 1986: 39–41, Fig. 2B. situation reminds us of what Natsukari Loligo yokoyae Ishikawa, 1926: 30–32, et al. (1975) reported concerning the Fig.1. misidentification of Loligo edulis as Lo- Loliolus rhomboidalis Burgess, 1967: 319– ligo chinensis by Wakiya & Ishikawa 329, Figs. 1–5, tab. 1. (1921), and it certainly supports the more Nipponololigo kobiensis, Natsukari, 1983: general statement by Okutani et al. (1987) 314–315. about the polymorphic characteristics of Nipponololigo sumatrensis, Natsukari, the species. 1984b: 262.—Dunning, 1998: 773. Again, ultimately the identification is Loligo sumatrensis d’Orbigny, 1839.— possible on the basis of the arm sucker Nesis 1987: 154, Fig. 34 g–i; 157, Fig. dentition, clearly edulis-type. While the 37 t–v. modified arms in males are hardly detect- Loliolus (Nipponololigo) sumatrensis, able, nevertheless the relatively very long Okutani, 1990: 59, Fig. 92.—Vecchione modified portion possesses the minute et al. 1998. suckers on the tips of the cone-shaped papillae. FAO vernacular names.—En—Kobe´ The above-mentioned differences in the squid; Fr—Calmar kobi; Sp—Calamar general appearance among our specimens kobi. from the different areas are not numeri- Type locality.—Japan cally quantifiable, nor even numerically Distribution.—Western Pacific (Map evident. Although quantitatively few, our 4), from southern Japanese waters south- observations tend to confirm the marked westward to Sumatra, including South polymorphism of the species already Korean waters, China, Thailand, Philip- emphasized by several authors in relation pines and Indonesia (Roper et al. 1984, to different geographical areas and, Okutani et al. 1987, Nesis, 1987), west- possibly, different seasons within the ward to the Bay of Bengal from the same area. Our results also underline the Andaman Sea to the east coast of India need for more detailed observations, and the Maldive Islands (Burgess 1967, information, and clarification on L. edulis Nesis, 1987, Nateewathana 1992, present throughout the Indian Ocean, especially work). because of its well-documented impor- Material examined.—Five specimens of tance as a fisheries resource in the western Loligo sumatrensis (3 males, 2 females) Pacific. from Madras, central east coast of India. 112 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

20* 40* 60" 00" 100" 120" 140*

40*

20* X 0* •

-20°

-40°

Map 4. Geographical distribution of Loligo sumatrensis, (shaded areas); open circle, present material, general locality.

See Appendix 1D for detailed localities lowish ground color with purplish red and capture data. All measurements and . These chromatophores relative indices are reported in Appendix are broadly distributed over the body 2D. Selected ranges of indices are re- surface without a distinctive pattern, ported in Table 8. except for a dark purplish midline dor- Diagnosis.—Squids of small size up to sally, in correspondence with the un- 120–130 mm ML (Fig. 4A, B). Fin length derlying gladius. generally about 60–65% ML; fin width The mantle is sub-cylindrical and grad- slightly larger at 60–70% ML. In males ually tapers posteriorly into a blunt point most of the left ventral arm is hectocoty- (slightly more pointed in females). Mantle lized, up to 87% (see Table 4A), and the width is about 30% of mantle length. Fins basal part of the right ventral arm bears are rhombic, rather wide (up to 70% of about 4 pairs of greatly enlarged suckers mantle length), with straight margins. Fin (Fig. 4G, H). Largest arm sucker rings length is about 55–57% of mantle length. with 6–9 large, very obtuse, square teeth, The head is small, with large eyes. Only almost as wide as high (Fig. 4F). Largest minor differences in general morpholog- medial manus sucker rings smooth ical features are noticeable between males (Fig. 4E). Smaller lateral manus sucker and females. The arms are moderately rings with acute teeth (Fig. 4D). long (slightly shorter in females), with an Description.—The material we exam- arm formula of 3.4.2.1. Several (up to 8) ined is in relatively good condition, and very enlarged globular suckers are present the general aspect conforms closely to the on the proximal end of the right ventral description given by Burgess (1967; the arm of males. Suckers of the other arms species, as L. rhomboidalis, is fully illus- are similar in shape and size in males and trated in this work based on Anton Bruun females, except the proximal suckers on IIOE material), including the coloration arms III of males are secondarily enlarged of the preserved specimens, a pale yel- at sexual maturity (Brakoniecki 1986). VOLUME 119, NUMBER 1 113

Fig. 4. Loligo sumatrensis d’Orbigny, 1835. A, ventral view (Roper et al. 1984; as Loligo kobiensis.B, dorsal view (redrawn from Burgess 1967). C, tentacular club (Roper et al. 1984). D, tentacular club, small toothed sucker ring, lateral row (Roper et al. 1984). E, tentacular club, large smooth sucker ring, medial row (Natsukari 1984). F, arm III sucker ring (Roper et al. 1984). G, hectocotylus, right arm IV (Nesis 1982/1987). H, hectocotylus, left arm IV (Nesis 1982/1987). I, gladius (Burgess 1967).

The left ventral arm is hectocotylized in crease in size distally; those of the ventral males; 6–7 normal suckers are present on series are more robust, palisade-like. The the proximal portion of the arm. The distal-most part of the arm remains un- modified portion follows, with the suck- modified in two thirds of the specimens in ers modified into two series of fleshy our study. papillae: those of the dorsal series are Tentacles weak, with clubs rather reduced, stump-like, and gradually de- strongly developed in comparison 114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Fig. 4C). Suckers of the median series Loligo singhalensis var. beryllae Robson, about 2–4 times larger than those on 1928: 15, Figs. 4–10. lateral series and flattened, with smooth, Doryteuthis singhalensis, Naef, 1912: 472. non-dentate rings. Suckers on the mar- Doryteuthis sibogae Adam,1954: 146, Fig. ginal series with small, acute teeth. 16–18, pl. I, Fig.4.—Silas 1986:31, The gladius is broad, paddle-like, with pl. VIII, A–B. broad, curved vane and long rachis Loligo sibogae, Natsukari, 1976: 15, Fig. (Fig. 4I). 1–14. A comparison of selected measure- Photololigo sibogae, Natsukari, 1984a: ments, indices and sucker dentition from 232.—Dunning, 1998: 777. various authors is given in Table 4C. Loligo (Photololigo) singhalensis, Okutani Biology.—Loligo sumatrensis is a neritic 1990: 61, Fig. 76. species that commonly inhabits coastal Uroteuthis (Photololigo) singhalensis,Vec- waters. Its biology has not been specifi- chione et al., 1998. cally investigated in this study, because of its marginal interest to fisheries. A FAO vernacular names.—En—Siboga synopsis of the available biological squid; Fr—Calmar siboga; Sp—Calamar knowledge, however, is given by Chotiya- siboga. putta (1993a), for the Gulf of Thailand Type locality.—Sri Lanka (Indian Ocean). and the Andaman Sea populations. Ac- Distribution.—Loligo singhalensis is an cordingly, L. sumatrensis is common in Indo-Pacific species. Its distribution (Map captures at sizes that range between 20 5) extends in the western Pacific from the and 70 mm ML, and its spawning period South China Sea [Singapore, Indonesian extends year round. waters by Adam (1954)], north to For- Fisheries.—Taken as by-catch in fish- mosa Island (Natsukari 1976) and west- eries for other squids in most of its ward into the Indian Ocean from the distributional area, e.g., southwestern Andaman Sea, Thailand (Nateewathana Japanese waters. It is highly abundant in 1992, Chantawong & Suksawat 1997); the the Gulf of Thailand and in the Andaman Bay of Bengal (Perera 1975, Silas et al. Sea (Chantawong & Suksawat 1997), 1985, Jothinagayam 1987), westward whereas it appears to be scarce in the (Dayratne 1993) to the Arabian Sea (Zuev Bay of Bengal. 1971) and the eastern African coasts Remarks.—Manoch (1998) hypothe- (Somalia by Rocha et al. 1998, present sised that Loligo uyii from the eastern work; Seychelles by Okutani 1970; Mo- side of the Gulf of Thailand is synony- zambique, at Mascarene underwater ridge mous with Loligo sumatrensis. by Pinchukov & Makarova, 1984). Material examined.—Twenty-four spec- Subgenus Doryteuthis Naef, 1912 imens of L. singhalensis (13 males, 11 Loligo (Doryteuthis) singhalensis females), from Djibuti, Somalia. See Ortmann, 1891 Appendix 1E for details of locality and [Uroteuthis (Photololigo) singhalensis, data collection. All measurements and Vecchione et al., 1998]. relative indices are reported in Appendix [Uroteuthis (subgenus undetermined) 2E. Selected ranges of indices are re- singhalensis, Vecchione et al. (2005)]. ported in Table 8. Fig. 5A–F Diagnosis.—Fin length in adults up to 50–60% of mantle length (Fig. 5A). Hec- Loligo singhalensis Ortmann, 1891: 676, tocotylized part of left ventral arm up to pl. 46, Fig. 3a–d.—Brakoniecki 1986: 45% of total arm length (Fig. 5E). Arm 25–26, Fig. 1F. suckers with 6 (5)–11 wide, squared, VOLUME 119, NUMBER 1 115

20* 40* ao* w 100* 120° 140°

40° V/\r **\ #^*^' 20°

20*

-40*

Map 5. Geographical distribution of Loligo singhalensis, (shaded areas); open circle, present material, general locality. plate-like teeth distally (Fig. 5D). Club cally described by Korzun & Alekseev suckers with 15–20–25 conical, sharply- (1991). pointed teeth around the entire margin, The head is comparatively small and that occasionally may be interposed by short; the eyes are of medium-size. smaller, almost inconspicuous, pointed The arms are slender and proportion- teeth (Fig. 5C). Gladius narrow, widest ally short. The arm formula varies: in its anterior 0.33, with nearly straight 3.2.4.1, as well as 3.4.2.1 (Ortmann margins on the vane (Fig. 5F). One 1891, Robson 1928, Adam 1954, Okutani photophore present on each side of the 1970, Natsukari 1976, Silas et al. 1985, ink sac. Jothinayagham 1987). The suckers are Description.—The mantle is long, slen- biserial, medium-sized, largest on arms II der, cylindrical, and it tapers posteriorly and III; sucker index 5 0.8–1.4 (Natsu- into a sharply-pointed tip. Mantle about kari 1976). The chitinous sucker rings 4–7 times as long as wide, widest at the are smooth or wavy proximally, while midpoint of its length; MWI around 19 in the distal margin bears 6–11 (most mature (males) (Table 5A). The commonly nine) plate-like, truncate, fins are narrow, rhombic in outline, with squared teeth. the anterior margin slightly convex and The left arm IV is hectocotylized in the posterior margin slightly concave. mature males for 40–45% of its length, by Fins are relatively short in young speci- the modification of suckers and stalks mens and become longer with growth along both rows into uniform, cone- (Natsukari 1976, Korzun & Alexeev shaped, pointed, fleshy papillae, devoid 1991), their length half or slightly more of minute suckers on their tips. The than half the length of the mantle in papillae on the ventral row are only adults. The relation between fin length slightly longer than those of the dorsal and mantle length has been mathemati- row. On the proximal part of the arm are 116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Loligo singhalensis Ortmann, 1891. A, dorsal view (FAO original). B, tentacular club (Nateewathana 1992). C, tentacular club, largest sucker ring (Roper et al. 1984; as Loligo sibogae). D, arm III, largest sucker ring (Roper et al. 1984; as L. sibogae). E, hectocotylus (Nateewathana 1992). F, gladius (Voss & Williamson 1972).

15–20 pairs of normal suckers. While the The tentacles are short and slender. hectocotylized portion of the arm be- Clubs are rather short (ClLI 5 12–25, comes relatively longer with maturity present work) (Fig. 5B). Club suckers (Natsukari 1976), the number of suckers quadriserial, those of the median two on the proximal part seems to remain rows of the manus slightly larger than constant. those of the lateral two rows. The largest VOLUME 119, NUMBER 1 117 suckers bear 15–20 incurved, conical, Vizhinjam coastal waters are 70–205 mm very sharp (5‘‘thorn-like’’, Ortmann ML in the fishery (Silas 1986). 1891), widely-spaced teeth around the Data from the southeastern slope of the entire chitinous ring. Occasionally a few Mascarene submarine ridge (Pinchukov smaller, almost inconspicuous and sharp, & Makarova 1984), western Indian secondary teeth are interposed between Ocean, indicate that this species is repre- the primary teeth: their presence is in- sented by two sympatric groups which dicated by the symbols (6) in Appendix differ slightly in size at first maturity. 2E. Males of the first group, which is The buccal membrane is well devel- widespread over the distributional oped, and each of its seven lappets bears area, reach sexual maturity at a size up to 12 small suckers. range of 80–210 mm ML and females Luminous organs are present, one on at a size range of 100–170 mm ML. each side of the ink sac. Based on this The second group occurs only in the character, the species should be included northern part of the area, where males in the proposed genus Photololigo (Nat- and females become mature at a size of sukari 1984a). 100-230 mm ML and 110–170 mm ML, The gladius is very narrow and slender, respectively. with almost straight lateral margins along Among the specimens examined in the the vane; it is widest at its anterior (0.33) present work, 6 males and 2 females were part. mature (46% and 18%, respectively, of Table 5A and C present the relative the total number available), with a ML indices obtained for this species in the range of 125–150 mm and 111–119 mm, present work, as well as those computed respectively; they were collected during by other authors. the month of February. Biology.—A semipelagic, neritic species Fishery.—Reported mainly as a by- that occurs mainly in coastal (inshore) catch in the Indonesian and South China waters, L. singhalensis has been reported Sea fin fisheries (as L. sibogae, Roper et to inhabit bottom depths down to 220 m al. 1984), L. singhalensis is likely to be (Pinchukov & Makarova 1984). The more important to the fisheries in Indian biology of this species is still poorly Ocean waters, even though the extent of known. Data from the southwest coast its contribution is difficult to assess of India (Silas 1986) indicate that males in because of confusion with the nomencla- that area attain sexual maturity at a size ture (see Remarks). For example, Chikuni range of 70–170 mm ML (50% mature at (1983) listed L. singhalensis among the 97 mm ML), and females mature at a size major species of loliginid squid that occur range of 70–130 mm ML (50% mature at in the Indo-Pacific region, with small 84 mm ML). All males and females are catches recorded from the China Sea to fully mature at a size of 170 and 130 mm, the eastern Arabian Sea, but it is not clear respectively. Mature specimens of both to what extent he actually referred to sexes occur from October to April, different species. suggesting a protracted spawning season. Included among the important species Juveniles are caught along the coast from for the commercial squid fishery of Thai- January–February up to June (in some land (Chantawong & Suksawat 1997), years), supporting this hypothesis. Based L. singhalensis seems consistently present on the analysis of the length frequency in the Andaman Sea (Chotiyaputta distributions, a life cycle of over two years 1993a), where it is one of the target has been estimated (Pinchukov & Makar- species of a light-attraction fishery in ova 1984, Silas 1986). Adults in the Phuket and Phang-Nga provinces (Chan- 118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tawong 1994). It is definitely among inclusion of L. singhalensis into the the important species for the Indian newly-created genus Photololigo of Nat- fishery (Ramachandran 1987). Reported sukari (1984a), and presented a summary as present on both coasts of the Indian of the situation concerning this species peninsula (Silas et al. 1982, as Doryteuthis in the literature, by indicating those sp. by Silas et al. 1985, Silas 1986, as references they consider valid for the Doryteuthis sibogae), it seems particularly species and those they do not consider important for the local fisheries off the valid, as follows: southwestern coast (Silas 1986). It also is likely to contribute consistently to Photololigo singhalensis the small pelagic local fishery in Sri (Ortmann 1891) Lankan waters (Dayaratne 1993, as Valid references: L. singhalensis). The species also probably Loligo singhalensis Ortmann, 1891. is important in the northern part of Loligo singhalensis var. beryllae Rob- the Bay of Bengal (Mohan & Rayudu son, 1928. 1986, as Doryteuthis sp.). However, it is Doryteuthis singhalensis, Naef, 1912.— not clear to what extent the references Okutani, 1970*.—Perera, 1975*.—Pin- to this species as the second most chukov & Makarova, 1979, 1984.— important squid resource of India in Zuev, 1971, Zuev & Nesis, 1971. general (Ramachandran 1987), and off the north-western coast in particular Doryteuthis sibogae Adam, 1954.— (Siraimeetan 1990), might include other Natsukari, 1976. species as well. Recent data (Meiyappan Loligo sibogae, Alekseev, 1989.—Nesis, & Mohamed, 2003) on Indian landings, 1982, 1987.—Roper et al. 1984. however, seem to confirm that L. singha- Photololigo sibogae, Natsukari, 1984. lensis (listed as Doryteuthis sibogae)isthe Invalid references: second most frequently caught squid, non Loligo singhalensis (5 L. chinen- representing 11% of east coast catches sis), Okutani, 1980.—Nesis, 1982, and about 2% of the catch from the 1987.—Alekseev, 1989. southwestern coast. non Doryteuthis singhalensis (5 L. Remarks.—The systematic position of edulis), Adam, 1939b, 1954. this species has been the object of several non Doryteuthis singhalensis (5 L. discussions and various contradictions chinensis), Voss, 1963.—Voss & Wil- during the past (Naef 1912: Adam 1939b, liamson, 1971;.—Roper et al.1984. 1954; Voss 1963: Silas 1968: Okutani 1970, non Photololigo singhalensis (5 L. 1980; Voss & Williamson 1971: Perera, chinensis), Natsukari, 1984. 1975: Natsukari 1976, 1984a; Nesis 1982, After our examination of the literature 1985; Silas et al. 1982, 1985: Silas 1986; we think it reasonable to agree with the Roper et al. 1984: Alekseev 1989, 1991; view of Korzun & Alekseev (1991). We Jothinayagam 1987), and of specific atten- also now refer to Loligo singhalensis the tion during the 1988 CIAC Workshop following references as valid: (Vecchione et al. 1998). Doryteuthis singhalensis, Jothinaya- Korzun & Alekseev (1991) gave a syn- gam, 1987*. thesis of the historical pathway of the Doryteuthis sp., Silas et al. 1982.— systematics of the species and came to Mohan & Rayudu, 1986. the conclusion that Doryteuthis sibogae Doryteuthis sibogae, Silas et al. 1985, Adam, 1954 is the earliest synonym Silas 1986. of Loligo singhalensis Ortmann, 1891. We do not refer to Loligo singhalensis Those authors did not object to the the following reference: VOLUME 119, NUMBER 1 119

Doryteuthis singhalensis, Silas et al. Silas et al. (1985) and Nateewathana 1986 (5 Loligo chinensis ?). (1992). Since our material agrees especial- In some cases (noted by*) the descrip- ly well with the description given by tion of some characters is ambiguous, i.e., Natsukari (1976), we suggest the reader the fin length is reported to be greater refer to this work for further details about than the range limits given for the species the species. by most authors; for example, Jothi- Genus Loliolus Steenstrup, 1856 nayagham (1987) reported ‘‘about 70%’’, Loliolus hardwickei (Gray 1849) Okutani (1970) measured ‘‘62%’’ and [Loliolus (Loliolus) hardwickei, Perera (1975) stated ‘‘…more than Vecchione et al. 1998]. half…’’ of mantle length, even though Fig. 6A–L in the latter case it is impossible to calculate a numerical FL/ML ratio. How- Loligo hardwickei Gray, 1849: 69. ever, the other characters defined and the Nesis, 1982: 132.—Lu et al. 1985: 70– illustrations of their specimens tend to 74, Fig. 7a–h, 8, tab. 3.—Nesis, 1987: support the identity of the subject species 146. with L. singhalensis Ortmann, 1891 Loliolus typus, Steenstrup, 1856: 194, (Jothinayagam 1987: 52, Fig. 18a, Oku- pl. 1, Figs. 5, 59.—Silas, 1968: 307. tani 1970, pl. 8 Fig. 4, Perera 1975: 54, Loliolus investigatoris Goodrich, 1896: 8– Fig. 6). 9, pl. 2, Figs. 29–37.—Massy, 1916: We consider the reference to Dory- 222.—Adam, 1939b: 66, Fig. 1.—Ad- teuthis sp. by Mohan & Rayudu (1986) as am, 1954: 29, Figs. 2–4.—Silas, 1968: valid for L. singhalensis, since these 307.—Oommen, 1975: 189.—Nesis, authors appear to adhere to the nomen- 1982: 132, Fig. 34 a,b,c,g.—Silas, clature used by Silas in numerous papers 1968: 307.—Silas, 1986: 21, 32, 33.— before 1985. Other species-specific refer- Silas, 1986: 116–118.—Jothinayagham, ences, mainly those reporting fishery 1987: 57–59, Fig. 20 a–g.—Nesis, 1987: statistics, unfortunately remain doubtful, 146. and their validation requires knowledge Loliolus hardwickei, Brakoniecki, 1986: about species-specific characters. 34–35, Fig. 2A. As for the ultimate assignment of this species to a genus and subgenus, it seems FAO vernacular names.—none. that the solution of this problem will Type locality.—India (Bay of Bengal ?). require some additional investigation. Distribution.—Loliolus hardwickei The revision accomplished by Vecchione (Map 6) is a common species throug- et al. (1998), who consider the correct hout the Indian Ocean, from the designation to be Uroteuthis (Photololigo) Persian Gulf, along the western and singhalensis, has been questioned again, eastern coasts of India, to the Andaman and following the CIAC meeting in Sea and into Indonesia waters (Silas Phuket (February 2003) the species now 1968, Silas 1986; Jothinayagam 1987, is considered to belong to the genus Nesis 1987). A single specimen was Uroteuthis, but to an undetermined sub- reported from the Chinese coast (Lu et genus (Vecchione et al. 2005). al. 1985). Following its original description (Ort- Material examined.—Forty-seven spe- mann 1891), this species was illustrated cimens of L. hardwickei (20 males, 27 by Naef (1912), Robson (1928) and females): 42 from the Arabian Sea and Adam (1954). More recently, exhaustive five from the Bay of Bengal, off Madras; and accurate descriptions have been given all Indian Ocean, mostly from the IIOE. by Okutani (1970), Natsukari (1976), See Appendix 1F for detailed localities 120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

20* 40* 00* 80* 100* 120* 140*

40*

20* f\ \ %. *% AL#= 0* ^

20*

-40*

Map 6. Geographical distribution of Loliolus hardwickei, (shaded areas); black dots, present material, site-specific localities; open circles, present material, general localities. and capture data. All measurements and general appearance and the main char- relative indices are reported in Appendix acters are almost exactly as described by 2F. Table 6A, B present general and Lu et al. (1985), enabling a confident regional measurements and indices; identification of the species. Table 6C provides a comparison of The mantle is short and stout (MWI up selected measurements, indices, and suck- to 58, Table 6A) with a conspicuous, er dentition from various authors. Select- rounded anterior dorsal lobe. Fins large, ed ranges of indices are reported in heart-shaped, with anterior and lateral Table 8. edges rounded; they occupy up to 76% Diagnosis.—Squid of small size, mantle of mantle length and their width is short, stout; fins posterior, broad, heart- greater than their length (FW 100–110% shaped, their width up to 100–110% of of ML). The fins continue past the mantle length (Table 6A; Fig. 6A–D). posterior end of the body, where they Left ventral arm hectocotylized (Fig. 6J). are fused. The modification extends along the Head short, a little narrower than the entire length of the arm (i.e., no normal mantle. suckers at the proximal base), and a pe- Arms rather short, sub-equal, usually culiar fleshy crest is present at the shorter in females than males. Arm proximal end. Arm suckers rings with 2– formula 3.4.2.1 (3.2.4.1 in type). 7 large, blunt, rounded, trapezoidal teeth Arm sucker rings with only 2–7 large, (Fig. 6H, I). Club sucker rings with 20– low, stout, broad teeth on the distal 40 small, sharp and/or truncate teeth margin; their number varies depending (Fig. 6F, G). on the arm and on the sex of the Description.—Most of the examined specimen. The proximal suckers on arms material is in a rather poor condition of II and III of males are enlarged as preservation (i.e., quite flabby), and some secondary sexual modifications (Brako- have already been dissected. However, the niecki 1986). VOLUME 119, NUMBER 1 121

Left ventral arm hectocotylized in (Table 6A; Lu et al. 1985). Maximum males by the modification of both ventral reported size is 88 mm ML for a male and dorsal series of suckers along its captured at the mouth of the River entire length: suckers are missing, except Hughli, India (Massy 1916). at the very tip where two to three minute Fishery.—In spite of its common suckers may be present. Ventral sucker occurrence in the Indo-Pacific region, stalks enlarged and fused entirely with the few data on catch statistics are avail- ventral trabeculate membrane, making it able. With so many other species of ‘‘…greatly enlarged and thickened into much larger loliginid squids in high a fleshy ridge.’’ (Lu et al. 1985:70). The abundance, probably not much demand proximal-most 2–3 trabeculae are further exists for such a small squid as L. enlarged (up to twice the membrane hardwickei. Perhaps artisanal fisheries height) to form a peculiar crest. Dorsal are developed in local situations. Loliolus series of sucker stalks reduced to small, hardwickei (as L. investigatoris)isre- free, lobate papillae. ported among the cephalopod resources Tentacles short. Clubs small (ClLI 15– of the northern part of the Arabian Sea, 40, Table 6A), scarcely expanded; i.e., the Guajarat coast (Siraimeetan 1990) manus club suckers sub-equal; suckers and the eastern coast of India (Silas of median series 1.0–1.3 times diameter 1986). of suckers of marginal series (Lu et al. Genus Sepioteuthis Blainville, 1824 1985) (Fig. 6E). Largest club sucker rings Sepioteuthis lessoniana Ferussac in with 20–40 small, acute or bluntly tri- Lesson, 1830 angular teeth around the entire margin. Fig. 7A–H Buccal membrane with seven-pointed lappets; each lappet bears 1–6 minute Sepioteuthis lessoniana Lesson, 1830: 244, suckers. pl. 11.—Sasaki, 1929: 127, pl. 14, Figs. Gladius broad (GWI 22-37, Lu et al. 15–17, pl. 29, Figs. 8–9, text Figs. 74– 1985); vane thin and fragile (Fig. 6K, L). 77.—Adam, 1939a: 21, pl. I, Figs. 1– Photophores absent. 2.—Voss, 1963: 77–81, Fig. 13.—Voss Additional details of the morphology & Williamson, 1971: 66–67, pl. 20, of L. hardwickei are presented in Lu et al. Figs. 19, 20, 26.—Lu & Tait, 1983: (1985). 183–190, Figs. 1–4, 8a.—Roper et al. Biology.—Very little information is 1984: 109–111.—Brakoniecki 1986: 15– available on the biology of L. hardwickei. 16, Fig. 1A.—Jothinayagam, 1987: 54– This species was not included with 57, Fig. 19.—Nesis, 1987: 146–147, Fig. the squids examined by Silas (1986), nor 35 i–m.—Okutani et al. 1987: 105.— was it in the resume´ of biological in- Nateewathana, 1992: 3–7, Fig. 1a–n, formation given for squids of the Anda- tab. 2.— Dunning 1998: 778.—Natee- man Sea by Chotiyaputta (1993a). Such wathana et al. 2000. limited attention undoubtedly is related Sepioteuthis hemprichii Ehrenberg, 1831: to the meagre importance of L. hard- [2]. wickei for the commercial fisheries, in Sepioteuthis guinensis Quoy & Gaimard, spite of its common occurrence in these 1832: 72, pl. 3, Figs. 1–7. areas. Sepioteuthis lunulata Quoy & Gaimard, Loliolus hardwickei is known to inhabit 1832: 74, pl. 3, Figs. 8–13. estuarine and coastal waters to a maxi- Sepioteuthis mauritiana Quoy & Gaimard, mum recorded depth of 30 m (Lu et al. 1832: 76, pl.4, Figs. 2–6. 1985). Males and females reach sexual Sepioteuthis sinensis Orbigny, 1848: maturity at a size of about 30–40 mm ML 304. 122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. A–G. Loliolus hardwickei (Gray 1849). A, dorsal view, female (Adam 1954). B, funnel notch on ventral mantle, female (Adam 1954). C, dorsal view, male (Adam 1954). D, funnel notch on ventral mantle, male (Lu et al. 1985). E, tentacular club (Lu et al. 1985). F, tentacular club sucker rings, female (Adam 1954). G, tentacular club sucker ring, male (Adam 1954). H–L. Loliolus hardwickei (Gray 1849). H, arm III sucker rings, female (Adam 1954). I, arm III sucker rings, male (Adam 1954). J, hectocotylus (Lu et al. 1985). K, gladius, female (Adam 1954). L, gladius, male (Adam 1954).

Sepioteuthis arctipinnis Gould, 1852: 479, Sepioteuthis neoguinaica Pfeffer, 1884: 4, Fig. 593. pl. I, Fig. 2, pl. II, Fig. 2a. Sepioteuthis brevis Owen, 1881: 137, Sepioteuthis indica Goodrich, 1896: 5, pl. 26, Fig.1. pl.1, Figs. 9–19. VOLUME 119, NUMBER 1 123

Fig. 6.—Continued.

Sepioteuthis sieboldi Joubin, 1898: 27. Distribution.—One of the most widely Sepioteuthis malayana Wu¨lker, 1913: 478, distributed loliginid squids (Map 7) of Fig. 7a–f. the Indo-West Pacific region, Sepioteuthis Sepioteuthis krempfi Robson, 1928: 28, lessoniana reportedly occurs from Figs. 3–4. Japan to Australia and New Zealand FAO vernacular names.—En—Bigfin and from Hawaii to the east African reef squid; Fr—Calmar tollelet; Sp— coast, north to the Red Sea and south Calamar manopla. to Madagascar (Adam 1939a, Chikuni Type locality.—‘‘Dorery.’’ Type not 1983, Roper et al. 1984, Nesis 1987, extant. Okutani et al. 1987, Rocha et al. 1998, 124 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Map 7. Geographical distribution of Sepioteuthis lessoniana, (shaded areas); black dots, present material, site-specific localities; open circles, present material, general localities.

B

Fig. 7. A–B. Sepioteuthis lessoniania Ferussac in Lesson, 1830. A, dorsal view, Indo-Pacific form (Roper et al. 1984). B. dorsal view, northeastern Australian form (Dunning and Lu 1998). C–H. Sepioteuthis lessoniana Ferussac in Lesson, 1830. C, tentacular club (Roper et al. 1984). D, tentacular club sucker ring (Roper et al. 1984). E, arm III sucker ring (Roper et al. 1984). F, hectocotylus, Indo-Pacific form (Roper et al. 1984). G, hectocotylus, northeastern Australian form (Dunning and Lu 1998). H, gladius (Voss & Williamson 1972). VOLUME 119, NUMBER 1 125

Fig. 7.—Continued.

Nateewathana et al. 2000, present mate- summary of selected measurements, in- rial). dices and sucker dentition from various Material examined.—Thirteen speci- authors is provided in Table 7C. Selected mens of S. lessoniana (5 females, 8 juve- ranges of indices are in Table 8. niles): 12 from East Africa, off Mombasa Diagnosis.—Fins broadly oval in out- and the western Madagascar coast, and 1 line, extend to the anterior edge of the from the Red Sea. See Appendix 1G for mantle, their length over 90% (up to detailed localities and capture data. All nearly 100%) of mantle length except in measurements and relative indices are paralarvae and small juveniles (Fig. 7A, reported in Appendix 2G. Table 7A gives B). Fins narrow in width uniformly from data on measurements and indices. A the point of maximum width, located at 126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON about the posterior 0.4 of the body (i.e., ical purposes. Consequently, the biology distance from widest point to the anterior of this squid has been studied rather edge of the mantle on average 60–65% of intensively, both in the field and in the mantle length). Each lappet of buccal laboratory (e.g., Rao 1954; Suzuki et al. membrane bears from 2–11 small suckers 1983; Silas 1986; Segawa 1987; Jackson (present material 4–6). 1989; Ueda & Jo 1989; Hanlon et al. Description.—A large loliginid squid 1991; Segawa et al. 1993a; Lee et al. 1994; (size exceeds 300 mm ML) with the Ueta & Segawa 1995; Ueta et al. 1995, Sepia-like body of the genus, unique 1999; Wada & Kobayashi 1995; Wada et among squids (Fig. 7A, B). The mantle al. 1995; Balgos & Pauly 1998; Boal & is long and robust, with the posterior end Gonzalez 1998; Ueta et al. 1999; Forsythe not sharply pointed, rather more bluntly et al. 2001; Mhitu et al. 2001; Pecl 2001; rounded. Fins very long, occupying al- Jackson & Moltschaniwskyj 2002; Samuel most the entire length of the mantle, their & Patterson 2002). width up to 75% (on average 60–65%)of Sepioteuthis squids, particularly S. se- mantle length. Head large with prominent pioidea of the western Atlantic, also are eyes. Arms strongly developed; arm known as ‘‘reef squids’’, due to their formula 3.4.2.1, the first pair smaller, preference for coral reef environments, more slender than pairs II–IV. Arm i.e., warm, clear, tropical and subtropical sucker rings with 18–29 triangular, sharp- waters (Vecchione et al. 1998). ly pointed teeth around the whole margin, Sepioteuthis lessoniana is a neritic spe- but most pronounced on distal 0.67 of cies that occurs from the surface down to ring (Fig. 7E). Tentacles long, robust; about 100 m depth throughout its area of tentacular clubs long (up to 35% of distribution. Rather common in coastal mantle length), only slightly expanded environments on sea grass beds, coral (Fig. 7C). Club sucker rings with 14-23 reefs, and sandy bottoms, it usually small, sharp, regularly spaced teeth migrates consistently inshore after the around entire ring (Fig. 7D). winter is over to start mating and spawn- Left ventral arm hectocotylized in ing (Rao 1954, Suzuki et al. 1983, Segawa males by the modification of the distal 1987, Segawa et al. 1993b, Ueta & one third (about 25–35%) of arm length, Segawa 1995). where the two series of suckers are Spawning season depends on the hy- transformed into long, conical, fleshy drographical conditions and can be quite papillae with a minute sucker with extended. It occurs from January to June smooth chitinous ring on each tip. Long, off southern India (Rao 1954) and from fleshy papillae connected by low folded mid-June to late August and September in ridge that extends along arm between the the cooler southern Japanese waters rows of papillae; papillae on dorsal row (Segawa 1987, Ueda & Jo 1989). Several thicker and longer than those on ventral peaks of two to three months during the row (Brakoniecki 1986) (Fig. 7F, G). year are reported for the Andaman Sea Gladius very wide, width about 20% of and the Gulf of Thailand populations length. Rachis stout, broad, short anteri- (Chotiyaputta 1993a). orly, gradually narrows posteriorly. The Recent observations confirm the exis- vane is slightly curved ventrally at its tence of flexible reproductive strategies in posterior end (Fig. 7H). this species (Pecl 2001) and affirm its Biology.—Sepioteuthis lessoniana is the potential to spawn multiple batches of subject of fisheries of various degree and eggs at discrete times throughout the adult importance, and it also is a very promis- life span. Therefore, even though field ing species for aquaculture and biomed- observations of spawned-out females do VOLUME 119, NUMBER 1 127 exist that show a consistent degeneration The employment of direct ageing tech- of the tissues of the whole body (e.g., Rao niques (i.e., statolith reading and analysis) 1954) and confirm that egg-laying is the on field-captured animals revealed that final event of the life cycle, the spawning S. lessoniana of the tropical Australian ‘‘phase’’ itself may be quite prolonged, as waters indeed do grow at a very fast rate. also indicated by rearing observations Individuals reached maturity and com- (e.g., Wada & Kobayashi 1995). pleted the whole life cycle in less than 100 Another example of the great flexibility days, at a size of about 180–200 mm ML of the S. lessoniana life cycle is given in (Jackson 1990). A very large (a the study of Jackson & Moltschaniwskyi mature male of 364 mm ML) was cap- (2002), who observed significant spatial tured and age-analysed by statolith at 164 and temporal variation in growth rates days (Jackson, pers. comm.), supporting and maturity of the equatorial, tropical, the hypothesis of a very rapid growth and subtropical Indo-Pacific populations. during the entire life cycle. In the ‘‘hot’’ shallow equatorial waters of The discrepancies between the results the Gulf of Thailand, S. lessoniana grow of length frequency analyses and statolith faster and mature earlier, at a much ageing were attributed, at least in part, to smaller size, than S. lessoniana that underestimation of the age of older squid inhabit the ‘‘cool’’ subtropical waters of (Balgos & Pauly 1998) in the latter case; southern Australia. The tropical popula- however, it must be remembered that tion off Townsville (northeastern Austra- length frequency analysis is not consid- lia) shows an intermediate situation be- ered to be an entirely adequate method to tween the two extremes, as well as an assess squid growth (e.g., Caddy 1991, alternation of generations that depends Alford & Jackson 1993, Jackson et al. on the season of hatching. 2000). Eggs are embedded in milky white, A rapid growth rate certainly is one of soft, gelatinous, slender, finger-like cap- the characteristics that makes S. lessoni- sules, each of which usually contains up ana especially suitable to laboratory to 13 eggs in a row (Segawa et al. 1993b). experiments and to aquaculture projects These capsules are attached in clusters to (Lee et al. 1994, Nabhitabhata 1995). the substrate. Based on the number of In addition, the life cycle in captivity is eggs in a single capsule, on the mode of completed in 4–6 mo, with adult weights capsule attachment to the substrate, and between 0.4 and 2.2 kg. Feeding rate is on the season of spawning, it was high; a variety of prey items, e.g., live suggested that S. lessoniana actually is fishes and crustaceans, is eaten, and late a complex of species in the Japanese juveniles and adults also can be trained to waters (Segawa et al. 1993a, Izuka et al. accept and ingest food pellets (Hanlon et 1994; see Remarks). al. 1991). Crowding in captivity can be Length frequency analyses and field a problem during the crucial juvenile observations indicate a life span that phase (Danakusumah 1999), but it is ranges between 1 and 3 years for this tolerated afterwards; the incidence of species (Rao 1954, Silas et al. 1982, disease and cannibalism is low. Repro- Segawa 1987, Ueda & Jo 1989). Rearing duction is easily achieved in captivity. experiments, however, evidenced a consid- All of these biological characteristics erably shorter life cycle, with females make S. lessoniana very interesting and reaching sexual maturity and spawning quite promising for future aquaculture at ages between 110 and 140 days projects focused not only on the needs of (ML between 105 and 145 mm, Segawa, biomedical research but also to those of 1987). human food production (Hanlon et al. 128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1991). Successful cephalopod mass-cul- lessoniana accounted for 7% of Indian ture experiments in Thailand on three east coast cephalopod landings, all from cephalopod species, including S. lessoni- Palk Bay and the Gulf of Mannar ana, have led to the production and (Meiyappan & Mohamed 2003). release of a consistent amount of cepha- Sepioteuthis lessoniana is reported as lopod paralarvae (about 2 million each moderately abundant in Sri Lankan year since 1990) to enhance natural stocks waters but rather scarce on the west coast (Nabhitabhata 1995). Recent information of India (Silas 1986). Very poor informa- reported that S. lessoniana has been tion is available for the remaining part of cultured through several successive gen- the Arabian Sea (Chikuni 1983) and erations (Walsh et al. 2002). eastern African waters, although we Fishery.—Sepioteuthis lessoniana is of expect that significant local fishery pro- commercial interest throughout its distri- duction occurs. butional range, being marketed mostly Remarks.—The systematic status of the fresh but also dried (e.g., Roper et al. genus has long been stable (Vecchione et 1984, Sarvaiya 1991). It is captured al. 1998), and since the generic revision by throughout the year with a variety of gear, Adam (1939a, four species are considered including lure-hooks, set nets, spears, valid, including S. lessoniana.Oneof beach seines or purse seines, and jigs in them, S. loliginiformis, is considered inshore waters (e.g., Dunning et al. 1994), doubtful by Nesis (1987). and by trawlers on the continental shelf. The hypothesis that two different infra- In Japanese waters, most squids are specific populations of S. lessoniana in- caught by directed trawl fishery and also habit the Okinawa area was postulated by as a by-catch, but a purse seine fishery Segawa et al. (1993a) on the basis of the directed at S. lessoniana occurs from May differences observed in the spawning sea- to September around Hong Kong (Chi- son, spawning substrate, attachment of kuni 1983); squid jigging also is efficiently capsules, number of eggs per capsule and practiced (Tokai & Ueta 1999). the arrangement of the deposited capsules The is one of the themselves. The ‘‘usual’’ egg capsules are loliginid species caught in the waters of found attached to an ‘‘exposed’’ substrate, the South China Sea (Nateewathana et al. contain a medium number of 4–8 eggs, and 2000), and it probably represents one of are deposited in April–May. The ‘‘un- the main components of the captures in usual’’ egg capsules are attached to the Indonesian waters (Chikuni 1983), as underside of hard substrata, contain only 2 evidenced by local studies (e.g., Soselisa eggs per capsule, and are laid from June to et al. 1986, Sudjoko 1987). It also is one October. Subsequent observations by iso- of the cephalopod species that supports zyme and other morphological analysis a minor domestic fishery in northern (Izuka et al. 1994, 1996a, 1996b; Yoko- Australian waters (Dunning et al. 1994). gawa & Ueta 2000) indicate the presence of This species is highly abundant in the three different taxa within the S. lessoniana Gulf of Thailand and in the Andaman complex off the Okinawa islands. They Sea, where it represents one of the most appear to be demonstrably reproductively important commercial species for the isolated; thus the species requires a system- cephalopod fishery (Chotiyaputta 1993a, atic revision. Chantawong & Suksawat 1997). It is caught in smaller quantities all along the Acknowledgments east coast of India (Silas 1986), and it supports a fishing industry in Madras The authors acknowledge with grati- state (Rao 1954). In recent years, S. tude the support of the Italian Consiglio VOLUME 119, NUMBER 1 129

Nazionale delle Ricerche (CNR) (Nation- ———. 1973. Cephalopoda from the Red Sea. al Research Council); the North Atlantic Contributions to the Knowledge of the Red Sea No 47.—Bulletin of the Sea-Fish Re- Treaty Organization (NATO); the Smith- search Station, Haifa 60:9–47. sonian U.S. National Museum of Natural Aleem, A. A., & S. A. Morcos. 1984. John Murray/ History, Department of Invertebrate Zo- Mabahiss Expedition versus the Interna- ology; the Smithsonian Oceanographic tional Indian Ocean Expedition (IIOE). Sorting Center, and the Rosensteil School In retrospect. Pp. 583–588 in M. V. Angel, of Marine and Atmospheric Sciences eds., Marine science of the north west Indian Ocean and adjacent waters 31 (RSMAS). We also acknowledge col- (6–8). leagues M. Vecchione (National Marine Alekseev, D. O. 1989. Advantages and limitations of Fishery Service Systematics Laboratory, using the gladius in diagnosis of species and Smithsonian U.S. National Museum of genera of the loliginid family.—Zoologi- Natural History) for helpful discussions chesky Zhurnal 68:36–42. ———. 1991. Systematics and phylogeny of bio- about loliginid systematics, and Nancy luminescent Loliginidae.—Bulletin of Marine Voss (RSMAS) for valuable comments Science 49:660. on the manuscript. The technical support Alford, R. A., & G. D. Jackson. 1993. Do of I. H. Roper is very much appreciated, cephalopods and larvae of other taxa grow as always. The comments and suggestions asymptotically?—American Naturalist 141: of two reviewers were most helpful and 717–728. Anderson, F. E. 1996. Preliminary cladistic analyses appreciated. of relationships among loliginid squids (Ceph- alopoda: ) based on morphological Note added in proof: data.—American Malacological Bulletin 12(1–2):113–128. While this paper was in press, an ———. 2000. Phylogenetic relationships among important volume was published that loliginid squids (Cephalopoda: Myopsida) addresses many aspects of loliginid based on analyses of multiple data sets.— systematics, life history, biology, and Zoological Journal of the Linnean Society fisheries. 130:603–633. Chotiyaputta, C., E.M.C. Hatfield and Ashirin, Z. S., & A. B. Ibrahim. 1992. Preliminary squid survey in coastal waters off Kuala C.-C. Lu, Guest Editors. 2005 Cephalo- Terengganu, Malaysia.—Asian Fisheries Sci- pod Biology, Recruitment and Culture; ence 5:261–264. International Symposium and Workshop, Asokan, P. K., & V. S. Kakati. 1991. Embryonic Phuket, Thailand, 2003. Phuket Marine development and hatching of Loligo duvauce- Biological Center Research Bulletin lii Orbigny (Loliginidae, Cephalopoda) in the laboratory.—Indian Journal of Fisheries 38: No. 66, 2005. 365 p. 201–206. Balgos, M. C., & D. Pauly. 1998. Age and growth of Literature Cited the squid Sepioteuthis lessoniana in N.W. Luzon, Philippines.—South African Journal Adam, W. 1939a. Cephalopoda, I. Le genre Sepio- of Marine Science 20:449–452. teuthis Blainville, 1824.—Siboga Expedition Bather, F. A. 1888. Shell-growth in Cephalopoda Monographie 55a:1–33. (Siphonopoda).—Annals and Magazine of ———. 1939b. The Cephalopoda in the Indian Natural History, series 6, 1. Museum, Calcutta.—Records of the Indian Berry, S. S. 1918. Report on the Cephalopoda Museum 41:61–100. obtained by the F.I.S. ‘‘Endeavour’’ in ———. 1954. Cephalopoda. Partie III. IV. the Great Australian Bight and other Cephalopodes a l’exclusion des genres Sepia, southern Australian localities.—Biological Sepiella et Sepioteuthis.—Siboga Expedition. Results of the Fishing Experiments carried Monographie 55(c):123–198. on by the F.I.S. ‘‘Endeavour’’ 1909-14 ———. 1962. Cephalopodes de l’Archipel du Cap- 4(5):201–298. Vert, de l’Angola et du Mozambique.— Blainville, H. M. D. de 1824. Mollusques, Memo´rias da Junta de Investigaco˜es do (Malacoz.).In F. Cuvier, ed., Dictionnaire des Ultramar, series 2, 33:9–64. Sciences Naturelles, Paris 32:1–392. 130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Boal, J. G., & S. A. Gonzalez. 1998. Social Chikuni, S. 1983. Cephalopod resources in the Indo- behaviour of individual oval squids (Cepha- Pacific region. Pp. 264–305, in J. F. Caddy, lopoda, Teuthoidea, Loliginidae, Sepioteuthis eds., Advances in assessment of world ceph- lessoniana) within a captive school.—Etholo- alopod resources, FAO Fisheries Technical gy 104:161–178. Paper 231. Boettger, C. R. 1952. Die Sta¨mme des Tierreichs in Chotiyaputta, C. 1982. Squid fisheries of Thailand. ihrer systematischen Gliederung.—Abhand- IPFC (Indo Pacific Fishery Commission) Re- lungen der Braunschweigischen Wissen- port of the Third Session of the Standing schlaftilchen Gesellschaft 4:238–300. Committee on Resources Research and Brakoniecki, T. F. 1986. A Generic Revision of Development. Sidney, Australia, 28 April– the Family Loliginidae (Cephalopoda; 4 May 1982.—FAO Fisheries Report Myopsida) Based Primarily on the 275:124–127. Comparative Morphology of the Hectocoty- ———. 1993a. Cephalopod resources of Thailand. lus. Unpublished Ph.D. dissertation, Uni- Pp. 71–80 in T. Okutani, R. O’Dor & versity of Miami, Coral Gables, Florida, 163 T. Kubodera, eds., Recent advances in pp. fisheries biology, Tokai University Press, Brock, J. 1887. Indische Cephalopoden.—Zoolo- Tokyo, 752 pp. gische Jahrbu¨cher 2:591–614. ———. 1993b. A survey on diversity and Brierley, A. S., A. L. Allcock, J. P. Thorpe, & distribution of juvenile squids in the R. D. Clarke. 1996. Biochemical genetic inner and western Gulf of Thailand.— evidence supporting the taxonomic Thai Marine Fisheries Research Bulletin separation of Loligo edulis and Loligo 4:19–36. chinensis (Cephalopoda: Teuthoidea) from ———. 1994. Distribution and abundance of the genus Loligo.—Marine Biology 127: juvenile and adult squids in the western Gulf 97–104. of Thailand. Pp. 200–207 in A. Snidvongs, W. Burgess, L. A. 1967. Loliolus rhomboidalis, a new Utoomprukporn & M. Hungspreugs, eds., species of loliginid squid from the Indian National Research Council Thailand–Japan Ocean.—Bulletin of Marine Science 17: Society for Promotion of Science Joint 319–329. Seminar on Marine Science, Chulalongkorn Caddy, J. F. 1991. Daily rings on squid statoliths: an University, Bangkok, Thailand, opportunity to standard population ———. 1997. Distribution, abundance, reproduc- models? Pp. 53–66 in P. Jereb, S. Ragonese tive biology, age and growth of Loligo & S. v. Boletzky, eds., Squid age determina- chinensis and Loligo duvaucelii in the western tion using statoliths. Proceedings of the Gulf of Thailand. Pp. 614–619 in D. A. International Workshop held in the Istituto Hancock, D. C. Smith, A. Grant & J. P. di Tecnologia della Pesca e del Pescato (ITPP- Beumer, eds., Developing and sustaining CNR), Mazara del Vallo, Italy, 9–14 October world fisheries resources, The State of Science 1989, N.T.R.-I.T.P.P. Special Publications 1, and Management, CSIRO, Collingwood, 137 pp. Australia, Chan, E. H., & A. K. B. M. Noor. 1986. Preliminary Cohen, A. C. 1976. The systematics and distribution data on squids (Loliginidae) from the of Loligo (Cephalopoda, Myopsida) in the Matahari Expedition. Pp. 221–227 in A. K. western North Atlantic, with descriptions of M. Mohsin, M. I. H. Mohamed & M. A. two new species.—Malacologia 15:299–367. Ambak, eds., Ekspedisi Matahari ’85. A Study Cuvier, G. 1797. Tableau e´le´mentaire de on the Offshore Waters of the Malaysian l’histoire naturelle des Animaux. Paris, 710 EEZ 3. pp. Chantawong, P. 1994. Squid light luring fishery in Danakusumah, E. 1999. Studies on the biology and Phuket and Phang-nga provinces, Andaman culture of the neritic squid Sepioteuthis Sea, Thailand.—National Research Council, lessoniana Lesson, 1830. Effects of stocking Thailand 93:184–194. density on survival rate. Pp. 223–226 in ———, & C. Suksawat. 1997. Cephalopod distri- J. Hylleberg, eds., Proceedings of the Ninth bution and abundance in the northern part of Workshop of the Tropical Marine Mollusc Phang-Nga Province, Thailand. Pp. 181–191 Programme, TMMP, Indonesia, 19–29 Au- in J. Hylleberg, eds., Proceedings of the gust 1998, Part 1 19(1). Seventh Workshop of the Tropical Marine Dayaratne, P. 1993. Exploitation of small pelagic Mollusc Programme, TMMP, on Central and fish resources by the purse seiners in the West Java, Indonesia conducted at IPB, southwest coast of Sri Lanka.—Journal of the UNDIP, LIPI, 11–22 November 1996, Part Marine Biological Association of India 1, 17(1). 35:39–45. VOLUME 119, NUMBER 1 131

Dunning, M. 1982. Squid and cuttlefish resources ———. 1886. Report on the Cephalopoda collected of Australian waters. IPFC (Indo-Pacific by H.M.S. ‘Challenger’ during the years Fishery Commission) Report of the Third 1873–1876.—Reports on the Scientific Re- Session of the Standing Committee on Re- sults of the Voyage of H. M. S. ‘Challenger’ sources Research and Development. Sidney, 1873–1876, Zoology 16, 246 pp. Australia, 28 April–4 May 1982. FAO Fisher- Ishikawa, C. 1926. Loligo yokoyae, a new species of ies Report 175:103–108. Myopsid Cephalopoda.—Proceedings of the ———. 1998. Loliginidae. Pp. 764–780 in K. E. Imperial Academy 2(1):30–32. Caprenter & V. H. Niem, eds., Living marine Izuka, T., S. Segawa, & T. Okutani. 1996a. Bio- resources of the western central Pacific, chemical study of the population heterogene- vol. 2, FAO, Rome, 1396 pp. ity and distribution of the oval squid Sepio- ———, S. McKinnon, C. C. Lu, & J. Yeatman. teuthis lessoniana complex in southwestern 1994. Demersal cephalopods of the Gulf of Japan.—American Malacological Bulletin Carpentaria, Australia.—Australian Journal 12(1–2):129–135. of Marine and Freshwater Resources 45: ———, ———, & ———. 1996b. Identification of 351–374. three species in oval squid, Sepioteuthis Ehrenberg, C. G. 1831. Cephalopoda in Mare lessoniana, complex by ar- Rubro viventia. In P. C. Hemprich & C. G. rangements on the funnel.—Venus, Japanese Ehrenberg, eds., Symbolae Physicae, seu Journal of Malacology 55:139–142. Icones et descriptiones Corporum Naura- ———, ———, ———, & K. Numachi. 1994. lium…quae ex itineribus per Libyan, Aegyp- Evidence of the existence of three species of tum…Habessiniam…Pars Zoologica, Berlin the Oval Squid Sepioteuthis lessoniana com- 4. plex in Ishigaki Island, Okinawa, Southwest- ern Japan, by isozyme analyses.—Venus, Fe´russac, A. E. de, & A. d’Orbigny. [1834] 1835– Japanese Journal of Malacology 53:217–228. 1848. Histoire naturelle ge´ne´rale et particu- Jackson, G. D. 1989. Age and growth of the tropical lie`re des Ce´phalopodes Ace´tabuliferes vivants nearshore loliginid squid Sepioteuthis lessoni- et Fossiles, lvi & 361pp; Atlas of 144 Plates. ana determined from statolith growth-ring Paris: J. B. Balliere. analysis.—Fishery Bulletin 88:113–118. Forsythe, J. W., L. S. Walsh, P. E. Turk, & P. G. ———. 1990. The use of tetracycline staining Lee. 2001. Impact of temperature on juvenile techniques to determine statolith growth ring growth and age at first egg-laying of the periodicity in the tropical loliginid squids Pacific reef squid Sepioteuthis lessoniana Loliolus noctiluca and Loligo chinensis.—The reared in captivity.—Marine Biology 138: Veliger 33:389–393. 103–112. ———. 1993. Seasonal variation in reproductive Goodrich, E. S. 1896. Report on a collection of investment in the tropical loliginid squid Cephalopoda from the Calcutta Museum.— Loligo chinensis and the small tropical sepioid Transactions of the Linnean Society of Idiosepius pygmaeus.—Fishery Bulletin 91: London, Series 2, Zoology 7:1–24. 260–270. Gould, A. A. 1852. Mollusca and shells. In United ———. 1995. Seasonal influences on statolith growth States exploring expedition … 1838–1842. in the tropical nearshore squid Loligo chinensis 12:1–510. (Cephalopoda: Loliginidae) off Townsville, Gray, J. E. 1849. Catalogue of the Mollusca in the North Queensland.—Fishery Bulletin 93: Collection of the British Museum, part I. 749–752. Cephalopoda Antepedia.—London, British ———, & N. A. Moltschaniwskyj. 2002. Spatial and Museum, 164 pp. temporal variation in growth rates and Guo, J., & P. Chen. 2000. A study on exploitation of maturity in the Indo-Pacific squid Sepio- Cephalopoda stock in South China Sea.— teuthis lessoniana (Cephalopoda: Loligini- Tropical Oceanology (Redai Haiyang Guang- dae).—Marine Biology 140:747–754. zhou) 19:51–58. ———, R. A. Alford, & H. J. Choat. 2000. Can Hanlon, R. T., P. E. Turk, & P. G. Lee. 1991. Squid length frequency analysis be used to determine and cuttlefish mariculture: an updated per- squid growth? An assessment of ELEFAN.— spective.—Journal of Cephalopod Biology ICES Journal of Marine Science 57:948–954. 2:31–40. Jasmine, G. I., C. B. T. Rajagopalasamy, & P. Hoyle, W. E. 1885. Diagnoses of new species of Jeyachandran. 1989. Total mercury content Cephalopoda collected during the Cruise of Indian squid Loligo duvaucelii Orbigny H.M.S. ‘Challenger’. Part II. The Deca- from Tuticorin waters, southeast coast of poda.—Annals and Magazine of Natural India.—Indian Journal of Marine Science History, series 5, 16:181–203. 18:219–220. 132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Jereb, P., & S. Ragonese. 1995. An outline of the Lamarck, J. B. 1798. Extrait d’un me´moire sur le biology of the squid Illex coindetii in the genre de la Se´che, du Calmar et Poulpe, Sicilian Channel (Central Mediterranean).— vulgairement nomme´s, Polypes de Mer.— Journal of the Marine Biological Association Bulletin des Sciences par la Socie´te´ Philoma- of the United Kingdom 75:373–390. tique de Paris 2(5):129–131. Jothinayagam, J. T. 1987. Cephalopoda of the Lee, P. G., P. E. Turk, W. T. Yang, & R. T. Hanlon. Madras Coast.—Zoological Survey of India, 1994. Biological characteristics and biomedi- Technical Monograph 15:1–85. cal applications of the squid Sepioteuthis Joubin, L. 1898. Sur quelques Ce´phalopodes du lessoniana cultured through multiple genera- Muse´e royale de Leyde et description de trois tions.—Biological Bulletin of the Marine esepces nouvelles.—Notes from the Leyden Biology Laboratory, Woods-Hole 186:328– Museum 20(2):21–28. 341. Karnik, N. S., & S. Kr. Chakraborty. 2001. Length- Lesson, R. P. 1830–1831. Mollusques, Anellides et weight relationship and morphometric study Vers. In Voyage autour du monde …sur la on the squid Loligo duvauceli (d’Orbigny) corvette de la Majeste, la Coquille, pendant (Mollusca: Cephalopoda) off Mumbai (Bom- les annees 1822–1825, Zoologie 2(1):239–471. bay) waters, west coast of India.—Indian Lesueur, C. A. 1821. Descriptions of several new Journal of Marine Sciences 30:261–263. species of Cuttle-Fish.—Journal of the Acad- Karnik, N. S., et al. 2003. Growth and mortality of emy of Natural Science of Philadelphia Indian squid, Loligo duvauceli (d’Orbigny) 2(1):86–101. (Mollusca: Cephalopoda: Teuthoidea) from Ling, J., & Y. Zheng. 2000. Stock assessment of Mumbai waters, India.—Indian Journal of Cephalopoda in East China Sea and Yellow Marine Sciences 32:67–70. Sea.—Marine Fisheries, Haiyang-Yuye, Kasim, H. M. 1985. Population dynamics of the Shanghai 2:60–62. squid Loligo duvaucelii d’Orbigny (Cephalo- Lu, C. C., & R. W. Tait. 1983. Taxonomic studies poda) in Saurashtra waters.—Journal of the on Sepioteuthis Blainville (Cephalopoda: Lo- Marine Biological Association of India liginidae) from the Australian region.—Pro- 27:103–112. ceedings of the Royal Society of Victoria Korzun, Yu. V. 1992. Finding of Loligo edulis 95:181–204. Hoyle, 1885, in the South-Western part of the ———, C. F. E. Roper, & R. W. Tait. 1985. A Indian Ocean.—Ehkol-Morya 41:66–68. [In revision of Loliolus (Cephalopoda: Loligini- Russian] dae) including L. noctiluca, a new species of Korzun, Yu. V., & D. O. Alekseev. 1991. Taxo- squid from Australian waters.—Proceedings nomic position of Loligo singhalensis (Cepha- of the Royal Society of Victoria 97:59–85. lopoda, Loliginidae).—Zoologichesky Zhur- Manoch, R. 1998. Re-examination of Loligo suma- nal 70:23–27. [In Russian] trensis d’Orbigny, 1835 along the eastern Kripa, V., et al. 1996. Cephalopod resources in coast of the Gulf of Thailand.—Thai Marine southeast and northeast coasts of India and Fisheries Research Bulletin 6:11–16. Andaman-Nicobar waters. Pp. 445–451 in V. ———, & F. Shunji. 1992. Some biological K. Pillai, S. A. H. Abidi, V. Ravindran & K. aspects of Indian squid, Loligo duvauceli, K. Balachandran, eds., Proceedings of the along the eastern coast of the Gulf of Second Workshop on Scientific Results of Thailand.—Thai Marine Fisheries Research Forv Sagar Sampada, New-Delhi, India De- Bulletin 3:45–53. partment of Ocean Development. Massy, A. L. 1916. The Cephalopoda of the Indian Kubota, T., T. Fukui, & T. Okutani. 1993. Museum.—Records of the Indian Museum Cephalopods incidentally caught with shirasu 12:185–247. boat seine from Suruga Bay in 1988, 1990 and Meiyappan, M. M., & K. S. Mohamed. 2003. 1991, with the catalogue of cephalopods Cephalopods. Pp. 221–227 in J. M. Mohan hitherto recorded from the bay.—Journal of & A. A. Jayaprakash, eds., Status of exploited the Faculty of Marine Science and Technol- marine fishery resources of India, Central ogy , Tokai University 35:77–91. Marine Fisheries Institutie, Kochi, India. Latif, M. S. A. 1982. Malaysian squid resources. Mhitu, H. A., Y. D. Mgaya, & M. A. K. Ngoile. IPFC (Indo-Pacific Fishery Commission) Re- 2001. Growth and reproduction of the big fin port of the Third Session of the Standing squid, Sepioteuthis lessoniana, in the coastal Committee on Resources Research and De- waters of Zanzibar. Pp. 289–300 in M. D. velopment. Sidney, Australia, 28 April–4 Richmond & J. Francis, eds., Marine science May 1982.—FAO Fisheries Report 175: development in Tanzania and Eastern Africa, 118–120. Zanzibar, Tanzania, 1. VOLUME 119, NUMBER 1 133

Mohamed, K. S. 1993. Spawning congregations of ———. 1991. Cuttlefish fisheries in Japan.—Acta of Indian squid Loligo duvaucelii (Cephalopoda: the 1st International Symposium on the Loliginidae) in the Arabian Sea off Manga- Cuttlefish Sepia, Centre de Publications de lore and Malpe.—Indian Journal of Marine l’Universite´e de Caen pp. 345–348. Science 22:172–175. ———, & T. Okutani. 1975. Taxonomic and ———. 1996. Estimates of growth, mortality and morphological studies on the loliginid squids. stock of the Indian squid Loligo duvaucelii I. Identity of Loligo chinensis Gray, 1849. Orbigny, exploited off Mangalore, Southwest Redescription of the type specimen and coast of India.—Bulletin of Marine Science taxonomic review.—Venus, Japanese Journal 58:393–403. of Malacology 34:85–91. ———, & G. S. Rao. 1997. Seasonal growth, stock- ———, & M. Tashiro. 1991. Neritic squid resources recruitment relationship and predictive yield and cuttlefish resources in Japan.—Marine of the Indian squid Loligo duvauceli (Orbigny) Behaviour Physiology 18:149–226. exploited off Karnataka coast.—Indian Jour- ———, T. Nakanose, & K. Oda. 1988. Age nal of Fisheries 44:319–329. and growth of the loliginid squid Photololigo Mohan, P. C., & G. V. Rayudu. 1986. Squid and edulis (Hoyle, 1885).—Journal of Experi- cuttlefish in nearshore bottom waters off mental Marine Biology and Ecology 116: Visakhapatnam, India. Pp. 373–377 in M. F. 177–190. Thompson, et al. eds., Biology of benthic ———, Y. Nishiyama, & Y. Nakanishi. 1986. A marine organisms, Balkema, Rotterdam. preliminary study on the isozymes of the Nabhitabhata, J. 1995. Mass culture of cephalopods loliginid Photololigo edulis (Hoyle, 1885).— in Thailand.—World Aquaculture 26:25–29. Report on the Cooperative Investigation of Naef, A. 1912. Teuthologische Notizen 1: Die ‘Shiro-ika’, Loligo edulis, Inhabiting the Familien der Myopsiden.—Zoologischer An- Western Japan Sea 2:145–151. zeiger 39:241–244. Nesis, K. N. 1982. [Abridged key to the cephalopod ———. 1916. Systematische u¨bersicht der Mediter- molluscs of the world’s ocean.], Light and ranean Cephalopoden.—Pubblicazioni della Food Industry Publishing House, Moscow, Stazione Zoologica di Napoli 1:11–19. 358 pp. [In Russian] Nateewathana, A. 1992. Taxonomic studies on ———. 1985. Oceanic cephalopods: Distribution, loliginid squids Cephalopoda:Loliginidae) life forms, evolution, Nauka, Moscow, 287 from the Andaman Sea coast of Thailand.— pp. Phuket Marine Biological Centre Research ———. 1987. Cephalopods of the world: squids, Bulletin 57:1–40. , octopuses and allies. Tropical ———, A. Munprasit, & P. Dithachey. 2000. Fish Hobbyist Publications, Inc., Neptune Systematics and distribution of oceanic ceph- City, New Jersey, 351pp. alopods in the South China Sea, Area 3: Western Philippines.—Proceedings of the Okada, Y. K. 1927. Cephalopodes japonais de Third Technical Seminar on Marine Fishery collections de Museum.—Bulletin du Muse- Resources Survey in the South China Sea, um d’Histoire Naturelle 93-98:172–179. Area 3: Western Philippines, 13–15 July 1999, Okutani, T. 1970. A small collection of gasteropod Bangkok, Thailand 41:76–100. and decapod mollusks from the Seychelles Natsukari, Y. 1976. Taxonomic and morphological Bank, Indian Ocean, by the training vessel studies on the loliginid squids. II. Descrip- Koyo-Maru in 1968.—Venus, Japanese Jour- tions and new records of Doryteuthis sibogae nal of Malacology 29:123–130. Adam, 1954, from Formosa.—Venus, Japa- ———. 1973. Guide and keys to squid in Japan.— nese Journal of Malacology 35:15–23. Bulletin of the Tokai Regional Fisheries ———. 1983. Taxonomical and morphological Research Laboratory 74:83–111. studies on the loliginid squid. III. Nippololigo, ———. 1975. Supplements and amendments for a new subgenus of the genus Loligo.—Venus, guide and keys to squid in Japan.—Bulletin of Japanese Journal of Malacology 42:313–318. the Tokai Regional Fisheries Research Lab- ———. 1984a. Taxonomical and morphological oratory 83:41–44. studies on the loliginid squid. IV. Two new ———. 1980. Useful and latent cuttlefish and squids genera of the family Loliginidae.—Venus, of the world. National Cooperative Associa- Japanese Journal of Malacology 43:229–239. tion of Squid Processors for the 15th Anni- ———. 1984b. Taxonomical and morphological versary of its Foundation, 66pp. studies on the loliginid squid. V. Re-de- ———. 1990. Cuttlefish and squid of the world in scription of the type specimen of Loligo colour. National Cooperative Association of sumatrensis d’Orbigny, 1835.—Venus, Japa- Squid Processors for the 30th Anniversary of nese Journal of Malacology 43:260–263. its Foundation, 185 pp, 50 pls. 134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

———, M. Tagawa, & H. Horikawa. 1987. shop of the Tropical Marine Mollusc Pro- Cephalopods from continental shelf and slope gramme (TMMP), 28 September–8 October around Japan. Japan Fisheries Resource 2000, 25. Conservation Association, 194pp. Quoy, J. R., & J. P. Gaimard. 1832. Mollusques. Oommen, P. 1975. New records of decapod In Voyage de de´couvertes de l’Astrolabe cephalopods from the Arabian Sea.—Journal pendant les annees 1826–1829, Zoologie of the Marine Biological Association of India 2(1):1–320. 17:188–190. Rahim, S. V. M. A., & M. R. Chandran. 1984. ———. 1976. Biology of the squid Loligo duvauceli Studies on the reproductive biology of the d’Orbigny obtained in the night catches.— warm water squid Loligo duvaucelii d’Or- Indian Journal of Marine Science 5:135–137. bigny. 1. Functional morphology of male D’ Orbigny, A. 1834–1847. Mollusques.—Voyage reproductive system.—Journal of the Marine dans l’Amerique Meridionale, 5(3):1–758, Biological Association of India 26(1–2): Atlas of 85 plates. 71–79. ———1845–1847. Mollusques vivants et fossiles ou Ramachandran, A. 1987. Cephalopod resources, description de toutes les espe`ces de Coquilles potential and utilisation in India.—Seafood et de Mollusques.605 pages, atlas of 36 plates. Export Journal 19:22–26. Paris. Rao, G. S. 1988. Biology of inshore squid Loligo Ortmann, A. 1891. Cephalopoden von Ceylon.— duvaucelii Orbigny, with a note on its fishery Zoologische Jahrbu¨cher 5:669–678. off Mangalore.—Indian Journal of Fisheries Owen, R. 1881. Descriptions of some new and rare 35:121–130. cephalopoda, part II.—Transactions of the Rao, K. V. 1954. Biology and fishery of the Palk- Zoological Society of London 11(5):131–170. bay squid, Sepioteuthis arctipinnis Gould.— Pecl, G. 2001. Flexible reproductive strategies in Indian Journal of Fisheries 1:37–66. tropical and temperate Sepioteuthis squids.— ———. 1973. Distribution pattern of the major Marine Biology 138:93–101. exploited marine fisheries resources of India. Perera, N. M. P. 1975. Taxonomic study of the Proceedings of the Symposium on Living cephalopods, particularly the Teuthoidea Resources of the Seas around India, Central (Squid) and Sepioidea (Cuttlefish) in the Marine Fisheries Research Institute, CMFRI, water around Sri Lanka.—Bulletin of the 1968.—CMFRI Special Publication 18–101. Fisheries Research Station, Sri Lanka Robson, G. C. 1928. Cephalopodes des mers 26:45–60. d’Indochine.—Service Oce´anographique des Pfeffer, G. 1884. Die Cephalopoden des Hamburger Peˆches de l’Indochine, Station Maritime de Naturhistorischen Museums.—Abhandlungen Cauda 10:1–53. aus dem Gebiete der Naturwissenschaften, Rocha, F., L. Fuentes, A. Guerra, & M. C. Sainza. Hamburg 8(1):1–30. 1998. Cephalopodos de Somalia.—Iberus Pinchukov, M. A., & L. M. Makarova. 1984. 16:129–142. Feeding and helminth fauna of the demersal ———, A. Guerra, & A. F. Gonzalez. 2001. A squid Doryteuthis singhalensis in the western review of reproductive strategies in cephalo- part of the Indian Ocean.—Malacological pods.—Biological Reviews 76:291–394. Review 17:130. Roper, C. F. E., & G. L. Voss. 1983. Guidelines for Prinngennies, D., & J. M. Jorgensen. 1994. Mor- taxonomic descriptions of cephalopod spe- phology of the luminous organ of the squid cies.—Memoirs of the National Museum of Loligo duvaucelii d’Orbigny, 1839.—Acta Victoria 44:48–63. Zoologica, Stokholm 75:305–309. ———, M. J. Sweeney, & C. E. Nauen. 1984. FAO ———, M. Murdjani, S. Sastrodihardjo, & N. R. species Catalogue, vol. 3. Cephalopods of the Nganro. 2000. A first attempt at culturing the world. An annotated and illustrated cata- squid Loligo duvaucelii Orbigny, 1835 in logue of species of interest to fisheries.—FAO Indonesia. Pp. 87–188 in J. Hylleberg, eds., Fisheries Synopsis 125, 277pp. Proceedings of the 10th International Con- Samuel, V. D., & J. Patterson. 2002. Intercapsular gress and Workshop of the Tropical Marine embryonic development of the big fin squid Mollusc Programme (TMMP), 20–30 Octo- Sepioteuthis lessoniana (Loliginidae).—Indian ber 1999, 21(1). Journal of Marine Sciences 31:150–152. ———, S. Sastrodihardjo, N. R. Nganro, & I. N. Sarvaiya, R. T. 1991. Prospect for export of dried Aryantha. 2001. Bacterial symbionts in squid from India.—Seafood Export Journal the luminous organ of the squid, Loligo 23:6–8. duvauceli, and cuttlefish, Sepia sp., Pp. 145– Sarvesan, R. 1974. Cephalopods.—Bulletin of the 148 in J. Hylleberg, eds., Proceedings of Central Marine Fisheries Research Institute, the 11th International Congress and Work- Cochin 25:63–83. VOLUME 119, NUMBER 1 135

Sasaki, M. 1914. Notes on the Japanese Myop- Song, H., T. Ding, K. Yu, J. Shen, X. Wang, & B. sida.—Annotationes Zoologicae Japonenses Ruan. 1999. Species composition and quan- 8(5):587–529. titative distribution of cephalopod in the ———. 1929. A monograph of the dibranchiate north of East China Sea.—Journal of the cephalopods of the Japanese and adjacent Zhejiang Ocean University of Natural Science waters.—Journal of the College of Agricul- 18:99–106. ture, Hokkaido Imperial University 20:1–357. Soselisa, J., S. Marjuki, & W. Subani. 1986. Segawa, S. 1987. Life history of the oval squid Production and fishing season of squids Sepioteuthis lessoniana in Kominato and (Loligo spp.) in Lombok (West Nusa Teng- adjacent waters of central Honshu, Japan.— gara) and adjacent waters.—Journal of Pene- Journal of the Tokyo University of Fisheries litian Perikanan Lautian [Journal of Marine 74:67–105. Fisheries Research] 34:79–90. ———, S. Hirayama, & T. Okutani. 1993. Is Steenstrup, J. 1856. Hectoctyldannelsen hos Octo- Sepioteuthis lessoniana in Okinawa a single podslaegterne Argonauta og Tremoctopus, species? Pp. 513–521 in T. Okutani, R. K. oplyst ved Iagttagelse af lignende Dannelser O’Dor & T. Kubodera, eds., Recent advances hos Blacksprutterne i Almindelighed.—Kon- in cephalopod fisheries biology, Tokai Uni- gelige Danske Videnskabernes Selskabs Skrif- versity Press, Tokyo, 752pp. ter, 5 Raekke, Naturvidenskabelig og Math- ———, T. Izuka, T. Tamashiro, & T. Okutani. ematisk 4:185–216. 1993. A note on mating and egg deposition by Sudjoko, B. 1987. Catch composition of squids Sepioteuthis lessoniana in Ishigaki Island, (Cephalopods) by using bagan (lift net) in Okinawa, southwestern Japan.—Venus, Jap- Probolinggo waters, East Java.—Journal of anese Journal of Malacology 52:101–108. Penelitian Perikanan Lautian [Journal of Shin, P. K. S. 1982. The Hong Kong squid Marine Fisheries Research] 41:81–89. fishery.—FAO Fisheries Report 175:112–114. Supongpan, M. 1984. The cephalopod fisheries and Shvetsova, V. D. 1974. Cephalopoda of the Indian resources in the Gulf of Thailand.—Thai Ocean.—Rybokhozyaistvennoe Ispol’zovanie Fisheries Gazette 37:340–346. Resursov Mirovogo Okeana, series 1, 8:4–18. ———. 1988. Assessment of Indian squid (Loligo Silas, E. G. 1968. Cephalopoda of the west coast of duvauceli) and mitre squid (L. chinensis)inthe India collected during the cruises of the Gulf of Thailand. Pp. 25–41 in S. C. Venema, research vessel ‘Varuna’, with a catalogue of J. M. Christensen & D. Pauly, eds., Con- the species known from the Indian Ocean. tributions to tropical fisheries biology, papers Proceedings of the Symposium on Mollusca, prepared by the participants at the FAO Part I.—Marine Biological Association of Danish follow-up training course on fish India, pp. 277–359. stock assessment in the tropics, Hhirtshals, ———. 1986 (ed.). Cephalopod bionomics, fisheries Denmark, 5–30 May 1986 and Manila, and resources of the exclusive economic zone Philippines, 12 January–6 February 1987, of India.—Bulletin of the Central Marine 389pp. Fisheries Research Institute 37, 195pp. Supongpan, M., & M. Sinoda. 1998. Sexual ———, K. P. Nair, & R. Sarvesan. 1985. New maturity size of Indian squid Loligo duvauceli record of a loliginid squid, Doryteuthis in the Gulf of Thailand.—Thai Marine sibogae, Adam 1954, (Cephalopoda; Loligini- Fisheries Research Bulletin 6:1–9. dae), from Indian waters.—Indian Journal of Suzuki, S., A. Kuwahara, & K. Washio. 1983. Study Fisheries 32:281–287. on the fishing conditions and an ecological ———, K. S. Rao, R. Sarvesan, K. P. Nair, & M. M. aspect of squids, Loligo edulis budo and Meiyappan. 1982. The exploited squid and Sepioteuthis lessoniana in the coastal waters cuttlefish resources of India: a review.—Ma- off Kyoto Prefecture.—Bulletin of the Japa- rine Fisheries Information Service, T & E Series nese Society of Fishery and Oceanography, 34, 16pp. Suisan Kaiyo Kenkyukaiho 42:21–27. Siraimeetan, P. 1990. Cephalopod resources of the Sweeney, M. J., & M. Vecchione. 1998. Generic and Gulf of Kutch, in Guajarat Coast.—Journal specific names introduced in the squid family of the Marine Biological Association of India Loliginidae (Cephalopoda: Myopsida).— 32:227–235. Smithsonian Contributions to Zoology 586: Sodikin, D. 1992. Review of squid fisheries in 223–237. Indonesia. Proceedings of the Second World Tokai, T., & Y. Ueta. 1999. Estimation of size Cephalopod Conference, Squid 991, Madrid, selectivity for oval squid Sepioteuthis lessoni- Spain, 11–13 November 1991. Agra-Europe, ana in the squid jigging fishery of Tokushima London, pp. 129–144. Prefecture.—Fishery Science 65:448–454. 136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Tryon, G. W. 1879. Manual of conchology. 1. ———, J. Nishioka, & M. Tanaka. 1995. On the Cephalopoda. Philadelphia, 316pp. spawning of the oval squid Sepioteuthis Ueda, Y., & Y. Jo. 1989. Notes on ecology of the lessoniana in the coastal waters off Kyoto oval squid Sepioteuthis lessoniana in outer Prefecture.—Nippon Suisan Gakkaishi 61: waters adjacent to the Kii Channel.—Nippon 838–842. Suisan Gakkaishi, Bulletin of the Japanese Wakiya, Y., & M. Ishikawa. 1921. Review of Society of Scientific Fishery 55:1699–1702. [In myopsid cephalopods in Japan with four Japanese]. new species.—Zoological Magazine, Tokyo ———, & S. Segawa. 1995. Reproductive ecology 33:279–292. and recruitment of juvenile of oval squid, Wallen, E., & H. A. Fehlmann. 1974. The Smithso- Sepioteuthis lessoniana in outer waters adja- nian oceanographic sorting center. A response cent to the Kii Channel.—Bulletin of the to the international Indian Ocean expedi- Japanese Society of Fishery and Oceanogra- tion.—Journal of the Marine Biological phy 4:409–415. [In Japanese]. Association of India 16:583–590. ———, I. Kitakado, S. Segawa, M. Tenjin, Y. Jo, Walsh, L. S., P. E. Turk, & P. G. Lee. 2002. M. Fukunaga, & Y. Kangawa. 1995. Selec- Mariculture of the loliginid squid Sepioteuthis tion of spawning ground and spawning bed in lessoniana through seven successive genera- oval squid Sepioteuthis lessoniana.—Suisan tions.—Aquaculture 212(1–4):245–262. Kogaku 31:189–194. [In Japanese]. Wooster, W. S. 1984. International studies on the Indian ———, T. Tokai, & S. Segawa. 1999. Relationship Ocean.—Deep Sea Research 31:589–598. between year-class abundance of the oval Wray, T. 1996. City feeds the world’s appetite for squid Sepioteuthis lessoniana and environ- squid.—Seafood International 11:41. mental factors off Tokushima Prefecture, Wulker, G. 1913. Cephalopoden der Aru- und Kei- Japan.—Fishery Science 65:424–431. Inseln.—Abhandlungen von der Senckenber- gischen Naturforschenden Gesellschaft 34: Vecchione, M., T. F. Brakoniecki, Y. Natsukari, & 451–488. R. T. Hanlon. 1998. A provisional generic Yeatman, J., & J. A. H. Benzie. 1994. Genetic classification of the Family Loliginidae. structure and distribution of Photololigo spp. Pp. pp. 215–222 in Voss, N. A., et al. eds., in Australia.—Marine Biology 118:79–87. Systematics and biogeography of cephalo- Yokogawa, K., & Y. Ueta. 2000. Genetic analysis of pods, vol. 1.—Smithsonian Contribution to oval squid (Sepioteuthis lessoniana)around Zoology 586:1–276. Japan.—Venus, Japanese Journal of Mala- ———, E. Shea, S. Bussarawit, F. Anderson, D. cology 59:45–55. Alexeyev, C.-C. Lu, T. Okutani, M. Roele- Zheng, Y. 1994. Present status and rational veld, C. Chotiyaputta, C. Roper, E. Jorgen- utilization on the exploitation of the resources sen, and N. Sukramongkol. 2005., Systematics of cephalopods in the East China Sea re- of Indo-West Pacific Loliginids.—Phuket gion.—Marine Fishery, Haiyang Yuye Marine Biological Center Research Bulletin 16:199–203. 66, 2005. ———, J. Ling, L. Yan, J. Zhou, & J. Shen. 1999. Vidyasagar, K., & V. D. Deshmukh. 1992. Stock Cephalopod resources and rational utilization assessment of Loligo duvaucelii (d’Orbigny) in in East China Sea.—Journal of Fishery Bombay waters.—Journal of the Marine Science, China 6:52–56. Biological Association of India 34:14–17. Zuev, G. V. 1971. Cephalopods of the Northwestern Voss, G. L. 1963. Cephalopods of the Philippines Indian Ocean.—Naukova Dumka, Kiev, Islands.—Bulletin of the United States Na- 223 pp. [In Russian] tional Museum 234:1–180. ———, & K. N. Nesis. 1971 (2003). The role of ———. 1973. Cephalopod resources of the world.— squids in the food chains of the ocean. FAO Fisheries Circular 149, 75pp. Chapter IV, Pp. 78–83. In Squid biology and ———, & G. Williamson. 1971. Cephalopods of fishing, 1971, Pishchevaya Promyshlennost, Hong Kong. Hong Kong Government Press Moscow, 360 pages. [In Russian]. English 138 pp, 35 pls. translation, 2003, Smithsonian Institution Wada, Y., & T. Kobayashi. 1995. On an iteroparity Libraries, 62–65. of oval squid Sepioteuthis lessoniana.—Nip- pon Suisan Gakkaishi 61:151–158. Associate Editor: Stephen L. Gardiner