Loliginidae LOLIGINIDAE Inshore Squids, Pencil Squids by M.C
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click for previous page 764 Cephalopods Loliginidae LOLIGINIDAE Inshore squids, pencil squids by M.C. Dunning iagnostic characters: Shape variable from short and stout to long and slender. Fins terminal or Dmarginal, but always reaching posterior end of body (present in Uroteuthis as very thin membranes around the “tail”). Funnel locking apparatus a simple, straight groove. Eyes covered with transparent membrane (cornea). Buccal connectives attached to ventral borders of arms (IV); 7 buccal lappets supplied with small suckers (can be readily lost after capture). Mouth surrounded by 10 appendages (8 arms, 2 tentacles). Two longitudinal rows of suckers on arms and 4 rows on tentacular clubs; hooks never present. Usually, left ventral arm (IV) hectocotylized in males; hectocotylus with suckers reduced in size or number, and/or modified into fleshy papillae or flaps, or lost completely. Colour: translucent to dense colour, bright scarlet (in some species with yellow and pink chromatophores) to dark brown (nearly black), darker dorsally, but highly variable depending on the behavioural situation and degree of expansion of chromatophores. ventral eye covered by funnel groove row of membrane suckers funnel dorsal row of suckers funnel pedicels locking cartilage modified mantle portion locking with cartilage reduced ventral (4th) suckers pair of arms hectocotylized corneal arm of males membrane gladius (internal skeletal mantle support) feather-shaped 4 longitudinal rows of suckers arms I beak (jaws) arm II buccal membrane tentacular club arm III buccal lappets often with suckers tentacle buccal connectives ventrally attached arms IV fin angle (left arm hectocotylized in males) ventral view diagram of oral surface of brachial crown and buccal area Loliginidae 765 Habitat, biology,and fisheries: Demersal or semipelagic inhabitants of coastal and continental shelf areas to a maximum depth of about 400 m. Several species are restricted to extremely shallow waters and some of these penetrate into brackish waters (Loliolus). Typically, they undertake diel movements, aggregating near the bottom during the day, but dispersing into the water column at night. Many species are positively phototactic, and hence often are captured with fishing techniques using light attraction. Some species undergo seasonal onshore-offshore migrations in response to temperature changes. The eggs (which range from about 2 mm diameter in Photololigo to about 10 mm diameter in Sepioteuthis) are encapsulated in gelatinous, finger-like strings and attached in clusters to various substrates. Hatchlings resemble the adults. Recent studies of growth using direct measurements from statoliths reveal that some tropical loliginids may live only for a few months. Inshore squids are predators on crustaceans and small fishes. Loliginids are the dominant component of the Western Central Pacific cephalopod catch, but separate statistics on total catches are not reported for this family. They are a significant component of major trawl fisheries in the Gulf of Thailand and South China Sea areas; they also are taken as bycatch in many nearshore trawl fisheries for crustaceans and demersal finfish. Numerous artisanal and subsistance fisheries take inshore squids either in multispecies catches or as prime target species; gear used includes purse seines, dip nets, lift nets, cast nets, encircling nets including fixed or tunnel nets in intertidal areas, baited and unbaited jigs sometimes trolled rather than used vertically, often used at night in association with light attraction. Loliginids are highly valued for human consumption. They are marketed either fresh, frozen, dried or processed into cleaned mantles (whole hoods, rings). Small squid are also used as bait for both commercial and recreational fisheries. Remarks: A recent study using modern biochemical genetic techniques has highlighted the incomplete state of knowledge of the taxonomy of the Indo-West Pacific loliginids. This is especially true for members of the genus Photololigo which includes the majority of the large commercially important species. The present set of diagnostic morphological characters (fin shape, sucker dentition, hectocotylus structure) does not always ensure a reliable identification of species. They do not take into account variability with growth and sex in some of these characters, particularly when these have been poorly and inadequately defined in the type descriptions (many from the mid 1800s). In many cases, type reference material is poorly preserved and had poor geographic locality information (e.g. Yokohama market for Loligo edulis, “Australian Seas (?S.E.)” for Loligo etheridgei. Generic level revisions have recently been undertaken for Sepioteuthis, Loliolus, and “Loligo”. However, additional studies at the species level over broad geographic areas are required to clarify the “species complexes” represented by what are currently known as “Photololigo edulis”, “P. c h i n e n s i s “ and ”Sepioteuthis lessoniana“. Several poorly known and new unnamed species are referred to in the literature and ”seasonal forms" with different life history characteristics and of questionable taxonomic status and distribution are known (e.g. Sepioteuthis lessoniana around Okinawa). This poor state of taxonomic knowledge has been highlighted previously in workshops and published papers but little progress has been made.Current and future fisheries assessments of the loliginid resource and subsequent decisions concerning the management of the stocks are dependent on accurate identification of species. Therefore, there is an urgent need for a substantial coopera- tive region-wide taxonomic study of the genus Photololigo using classical morphology supported by modern techniques including allozyme electrophoresis and DNA analysis. Similar families occurring in the area Sepiidae (cuttlefishes): the loliginid genus Sepioteuthis shows some external similarities to cuttlefish but can be easily distinguished from them by the presence of a gladius in the dorsal mantle rather than a chalky cuttlebone with a posterior calcareous spine; the presence of the midline con- nection of the fin lobes posteriorly in Sepioteuthis and the possession of contractile rather than retractile tentacles only without pockets. Identification note The key and descriptions provided below are based on the calcareous existing, inadequate adult character sets and users should spine recognize that in some cases, they may not readily assist Sepiidae Loliginidae (Sepioteuthis) with identifying the animals at hand. It should also be noted that in some cases, known or suspected species complexes are referred to rather than single species (“Photololigo chinensis”, “Photololigo edulis” and “Sepioteuthis lessoniana“). Some nominal species described in the literature and listed below are not included in the key because detailed descriptions have yet to be published, they remain poorly characterized from the type descriptions or are described from limited material and their status as valid species remains questionable. These include: 766 Cephalopods Loligo reesi (Voss, 1962), a form that matures at a small size (less than 80 mm mantle length) known only from the Philippines which has long square-tipped arm sucker teeth, both ventral arms hectocotylized in males (left with papillae along more than 50% arm length and right with much reduced suckers on the distal half) and the the tips of both ventral arms in males devoid of suckers and papillae. The existing descriptions do not mention the presence of light organs; Loligo vossi Nesis, 1982 [= species A of Voss (1963)], reaching 140 mm mantle length and reported from around the Philippines and northern Indian Ocean perifery, with 15 to 20 triangular teeth on the large tentacular suckers, 2 to 4 on the distal edge much larger, sharp and hook-like in Philippine specimens. Only the left ventral arm is hectocotylized in males by the development of long fleshy papillae along 40 to 50% of the arm. The existing descriptions do not mention the presence of light organs; Loligo n.sp. of Chotiyaputta (1993), a form that matures at a small size (less than 60 mm mantle length) known only from the Gulf of Thailand which has long square-tipped arm sucker teeth and sharp conical widely spaced subequal teeth on the large tentacular sucker rings. The existing incomplete description does not mention the presence of light organs; Photololigo spp. 1, 2, 3 of Yeatman and Benzie, 1994, are “species” which have been distinguished using allozyme electrophoresis. Morphological characteristics of species 1 and 2 from northern Australia (which mature at less than 120 mm mantle length) fall within the range of the “Photololigo edulis” complex with square tipped teeth on the arm sucker rings, sharp conical teeth on the large tentacular sucker rings and hectocotylization by development of long fleshy papillae along more than 50% of the left ventral arm in males. Species 3 is a larger maturing form (more than 150 mm mantle length) which falls within the “Photololigo chinensis” complex with a broad, robust mantle and large head and arms; sharp conical toothed sucker rings on the arms and large sharp conical teeth interspersed with smaller ones on the large tentacular sucker rings. All species have paired light organs adjacent to the rectum on the ink sac. It is recommended that samples of animals (including mature males and females where available) which cannot be identified readily from the key should be preserved in formalin or alcohol,