Occurrence of Idiosepius (Mollusca: Cephalopoda) in Indonesian Waters

Jurnal Biologi Indonesia 6 (1):13-23 (2009)

Occurrence of Idiosepius (Mollusca: Cephalopoda) in Indonesian waters

Janek von Byern 1 & Ristiyanti M. Marwoto 2 1 University of Vienna, Fac. of Life Science, Cell Imaging and Ultrastructure Research, 1090 Vienna, Austria 2 Museum Zoology Bogor, Research Center for Biology – LIPI, Cibinong 16911 – Indonesia;email: [email protected]

ABSTRAK

Jenis Idiosepius (Mollusca: Cephalopoda) di perairan Indonesia. Informasi dan penelitian tentang sotong mini “pygmy squid” marga Idiosepius yang ada di Indonesia sangat kurang, meskipun pernah dilaporkan setidaknya ada tiga jenis dijumpai di Ambon, Ternate, Banda, Balikpapan, Sibolga dan Lombok. Dalam tulisan ini diuraikan karakter morfologi, habitat dan distribusi empat jenis sotong mini jenis I. picteti, I. pygmaeus, I. biserialis dan I. pygmaeus herbereri. Karena ukurannya yang sangat kecil, jenis sotong ini tidak diminati oleh para nelayan, sehingga hanya sedikit data yang diketahui mengenai pertumbuhan, reproduksi dan siklus hidupnya. Penelitian ini bertujuan untuk memberikan gambaran ringkas tentang sistematik, habitat, siklus hidup dan distribusi Idiosepius. Hasil studi ini mencatat lokasi baru (new record) ditemukannya sotong mini jenis I. biserialis dan I. pygmaeus khusus dari perairan pantai di Lombok. Hasil penelitian juga menunjukkan bahwa beberapa jenis Idiosepius memiliki sebaran yang luas di perairan Indonesia kecuali jenis I. picteti hingga saat ini hanya dijumpai di Ambon. Hasil studi diharapakan menambah khasanah pengetahuan tentang sotong mungil ini sekaligus memacu para peneliti untuk lebih memperhatikan genus ini.

Keywords: Cephalopoda, distribution, habitat, Idiosepiidae, Indo-Pacific Kata kunci: Cephalopoda, distribusi, habitat, Idiosepiidae, Indo-Pacific

Introduction The animals are dorso-ventrally compressed and cigar-shaped; the fins The genus Idiosepius has the are small, ovally elongated with the long smallest species within the cephalopods axis parallel to the body axis (Figure 1). and is also named “pygmy squid”. The The arms are short and robust and almost females reach maturity at 3 cm length, equal in length, except for one arm that whereas the males of some species is always shorter (Hylleberg & reach sexual maturity at < 1 cm (Joubin Nateewathana 1991b). Both sexes can 1902; Norman 2000) with a body weight be distinguished easily by the modified ranging from 6 mg (Idiosepius fourth arm pair in males (Yamamoto biserialis) to 1 g (Idiosepius pygma- 1949; Jackson 1988; Nabhitabhata 1998; eus) (Hylleberg & Nateewathana Norman 2000). 1991a; Hylleberg & Nateewathana One conspicuous morphological 1991b). character of this genus is the adhesive

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Figure 1: Individual of the species Idiosepius pygmaeus. The animals are small in size and live exclusively in mangrove areas in the Indo-Pacific area down to Australia. organ (also named adhesive gland) and Banda-Sea. This species was located on the posterior part of the dorsal thought to occur only in the Philippines. mantle side (von Byern et al. 2008; Later, collections by Grimpe in 1931 in Cyran et al. 2008). The animals use the Balikpappan and Sibolga showed that I. glue from the adhesive glands to stick to pygmaeus has a wide distribution in sea grass leaves or algae for camouflage Indonesia. when threatened by predators (Sasaki In 1927, Rensch collected 14 1921). Hiding there, they also lie in wait specimens of the genus Idiosepius at to capture prey swimming by. Further- Ekas Bay, Lombok, Indonesia. Grimpe more, in females, secretion of glue (1931) compared the collected material serves to stick the eggs on sea grass with samples of I. pygmaeus from (Nesis 1982). several institutions and proposed that his Collection of Idiosepius in Indonesian specimens differ from I. pygmaeus and waters has long history; more than 115 the other species mostly in size, contour years ago, Joubin (1894a) collected one of the adhesive organ and expression of male holotype sample of I. picteti at the hectocotylus (Grimpe 1931). He Ambon Island. Unfortunately, up to this therefore regarded his specimens as the day, attempts to re-collect individuals of subspecies Idiosepius pygmaeus this species in its original type locations hebereri. or neighbouring island were unsuccess- Examinations by Nesis (1982), ful. It can be assumed that this species however, revealed that the morphological has disappeared or even become extinct. characteristics were insufficiently In 1898, Appellöf discovered I. different to justify the subspecies; pygmaeus Steenstrup, 1881 in Ternate Idiosepius pygmaeus hebereri was

14 Occurrence of Idiosepius (Mollusca: Cephalopoda) therefore referred to I. pygmaeus. In ches Museum Wien, Austria) and MZB contrast to I. picteti, individuals of I. (Museum Zoologicum Bogoriense), pygmaeus could still be located at their Research Center for Biology – LIPI, type locality (von Byern & Klepal 2007) Indonesia. as well as in new localities (see description of I. pygmaeus below). RESULT Apart from I. picteti and I. pygmaeus, a further species, I. Description of Indonesian biserialis, was recently discovered in Idiosepius species Indonesian waters (von Byern et al. Idiosepius biserialis Voss, 1962 (Figure 2005). So far, less is known about the 2A) geographical distribution and habitat of this species (see section Habitat Holotype: conditions in Indonesia). Previously, it deposited at South African Museum, was thought to occur only in African Cape Town, South Africa (SAM A6520) waters (Mozambique) and Thailand. With the present description and Type locality: geographical data, we report new Mozambique, San Jose Mission Station, collection places for Idiosepius in Morrumbene Indonesia and extend the previous type localities. This will shed light on the Geographical distribution of the ecology and distribution of Idiosepius species or /and material examined: and yield new insights into the habitat Thailand: Bang Rong, Phuket Island (8° conditions of Indonesian individuals. 02.156Ń; 98° 25.487´ E) (von Byern Nevertheless, many questions about & Klepal 2009) and Ko Pratong, their life cycle, geographical distribution Ranong (Hylleberg & Nateewathana and origin of migration remain. Further 1991a) collections coupled with ecological and Mozambique: Inhaca Island (26° behavioral investigations are necessary 00.215´S, 32° 54.721É; 26° 02.300´S, to complete our picture of this genus. 32° 54.166É) (Kalk, 1959; von Byern & Klepal 2009); Inhambane Bay (23° METHODS 51.184´S, 35° 22.553É) (von Byern & Klepal 2009); Monque (23° Idiosepius specimens were 41.331´S, 35° 22.281É) (Voss 1962; collected along the bay of Lombok in Adam 1986; von Byern & Klepal November - December 2007 using a 2009) dipnet. The specimens were then Japan: Takasu, Japan. Collector: S. preserved with ethanol 95% (partly for Shigeno (von Byern et al. 2005) DNA) and 70 % (for ordinary preserved Indonesia: Ekas-Bay, Lombok (08° collections). The specimens are 52.020´S; 116° 27.541É) (von Byern deposited at the NMW (Naturhistoris- et al. 2005) ; Ekas-Bay, Lombok (08°

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50. 357‘S ; 116° 27.857‘ E) (08° 50. in size to the end of the club. Both ventral 714‘S ; 116° 26.781‘ E) , collected arms in the male are hectocotylized, November 2007 (unpubl. data) bearing 1-5 suckers on the left (ventral view) and 3-8 suckers on the right ventral Morphological characteristics: arm. The left arm also has two small This species has the smallest specimens flaps, separated by a deep cleft, at the of the genus. Animals from Mozambique tip. In the caught male, the hectocotyli are small, slender, with a mantle length arms are almost unequal in length: the of 4.5 ± 0.3 mm (males) and 7.38 ± 1.31 left arm is shorter than the right one. mm (females). The fins are semicircular The specimens from Thailand are and about ¼ of the mantle length. The larger (ML to 5.84 ± 0.54 mm males and clubs of this species bear two rows of 7.7 ± 1.1 mm females). The fins are small suckers, which are nearly constant more rounded kidney-shaped and

14.2 mm

7.35 mm 15.3 mm

Figure 2: Schematic drawing of the known Idiosepius species in Indonesian waters: A) I. biserialis (Voss, 1962); B) I. picteti (Joubin, 1894a) and C) dorsal and ventral view of I. pygmaeus (Nesis, 1982).

16 Occurrence of Idiosepius (Mollusca: Cephalopoda) attached to the body at an angle. The ventral arm in the male is very short and tentacle clubs have two horizontal rows broad. The oral face the arm is of suckers, sometimes also three or four transversely plicate. The left ventral arm oblique rows of suckers. The hectoco- is more slender, longer and bilobate at tylized arms also bear 2-4/2-5 suckers the tip. Each ventral arm has a single on the left/right ventral arm. The animals small sucker near the base. Joubin from Japan are almost twice as long (1894b) wrote that the examined (6.34 ± 0.89 mm males and 9.39 ± 1.1 specimen has no adhesive organ on its mm females) with 3 suckers on the left dorsal side: “Pas dímpression dorsale and 5 on the right ventral arm. The entre les nageoires”. Our examinations specimens from Indonesia have a mantle of this specimen reveal that Joubin length of 3 mm (and 4 suckers on each (1894a) erred. Idiosepius picteti has an hectocotylized arm) in the one available adhesive organ on the dorsal mantle side male, while the two investigated females and is referred to the genus Idiosepius. have a mantle length of 5.25 ± 0.35 mm. Idiosepius pygmaeus Steenstrup, Idiosepius picteti Joubin, 1894b 1881 (Figure 2C). (Figure 2B) Synonymy: Synonymy: Idiosepion pygmaeum Fischer, 1887 Loligo picteti Grimpe, 1920 Idiosepius pygmaeus Hoyle, 1886; Naefidium picteti Grimpe, 1920 Joubin 1894b; Appellöf 1898 Idiosepius pygmaeus pygmaeus Holotype: deposited at the Muséum Grimpe, 1931 d’Histoire Naturelle, Genève, Swit- zerland (MHNG M 3/75 747/27) Holotype: deposited at Kobenhavns Universitet, Zoologisk Museum, Type locality: Indonesia, Amboina – Copenhagen, Denmark (ZMUC CEP- according to Joubin, 1894b (today Ambon 52) Island) Type locality: Philippines, Zamboanga Geographical distribution: only (4° 20Ń, 107° 20É) known from the type locality Geographical distribution of the Morphological characteristics: So far species or/and material examined: only the holotype is available; no Philippines: Jolo Habor (Adam 1986) additional specimens of this species have Thailand: Klong Mudong (7° 48.107Ń, ever been found. The body is large and 98° 24.472É) (von Byern a & Klepal elongate (mantle length 14 mm). 2009), Klong Bang Rong (8° Idiosepius picteti has a small tentacle 02.945Ń; 98° 25.030É) (Suwanmala club with four rows of suckers. The right

17 Byern & Marwoto

et al. 2006) and Ao Chalong right ventral arm is stout and thick, while (Hylleberg & Nateewathana 1991b) the left arm is thinner and slender, Singapore: Tempenisi (Adam 1986) bilobated at the tip. According to the Australia: Townsville, North Queensland description of Steenstrup (1881), both (19° 15´S, 146° 50´E) (Jackson 1988; ventral arms have only one sucker at Lewis 1991; Jackson 1992; Jackson their base. Later examination of this & Choat 1992; Jackson 1993; species revealed more numerous suckers Semmens et al. 1995; Pecl & in different variations on the ventral arms Moltschaniwskyj 1997; Pecl & (Appellöf 1898). The range of sucker Moltschaniwskyj 1999) combinations varies from 0 to 4 suckers Micronesia: Palau (Belau) Islands on the hectocotylized arms (Thailand). (Moynihan 1983) More than 30% of the specimens bear Indonesia: Ternate (Appellöf 1898); 2/3 suckers on the left/right or 3 suckers Balikpappan (Grimpe 1931); Sibolga on both ventral arms. (Grimpe 1931) and Banda-Sea Idiosepius pygmaeus from Indonesia (Appellöf 1898) is similar in size (11.53 ± 1.28 mm males New localities in Indonesia and 15.5 ± 2.12 mm females) to I. • Ekas-Bay, Lombok (08° 52.020´S; pygmaeus from Thailand and bears 2 or 116° 27.541´E) (von Byern & Klepal 3 suckers on the left and 3 suckers on 2007) the right ventral arm; one individual has · Gili Sulat (08° 19.885‘ S; 116° 43.015‘ 1/1, another 4/4 suckers on the E), collected November 2007 hectocotyli. (unpubl. data) • Gili Lawang (08° 19.637‘ S; 116° I. pygmaeus (= Idiosepius pygmaeus 42.709‘ E), collected December hebereri) Grimpe, 1931 2007 (unpubl. data) • Telong Elong (08° 48.730‘ S; 116° Holotype: deposited at the 30.012‘ E), collected December Zoologisches Museum, Museum für 2007 (unpubl. data) Naturkunde der Humboldt-Universität, • Rinca Island (08° 39.244‘ S; 119° Institut für Systematische Zoologie, 42.937‘ E), collected December Berlin, Germany (ZMB) 2007 (unpubl. data) Type locality: Indonesia, Ekas-Bay, Morphological characteristics: Lombok The species has a sepiolid body shape with a mean mantle length of 11.51 Geographical distribution: ± 1.52 mm in males and 16.26 ± 4.37 only known from the type locality mm in females. The fins are small, short, rounded and slightly constricted at their Morphological characteristics: base. The specimens bear 4 rows of Because of the close morphology to suckers at the club of the tentacle. The Idiosepius pygmaeus, Grimpe (1931)

18 Occurrence of Idiosepius (Mollusca: Cephalopoda) subordinated this species as Idiosepius (Nesis et al. 2002). Within the genus, I. pygmaeus hebereri. The females of this biserialis has the widest geographical subspecies have a mean mantle length distribution, ranging from Japan to the of 14.5 mm, the males of 8.5 mm. The Indo-Pacific (Thailand and Indonesia) fins are small and have a round to almost (Voss 1963; von Byern et al. 2005); it oval form. Grimpe (1931) did not has also been recorded in Mozambique describe the number of rows on the club (incorrectly annotated by Voss in 1962 but later re-examination of the holotype as South Africa). material showed that the specimens have four rows of suckers (von Byern, unpubl. Habitat and life cycle data). The left ventral arm is longer and The habitat occurrence varies within bigger than the right arm, and also has the genus: some species occur in two lobes at its tip. At its base the left mangrove areas (e.g. I. pygmaeus). arm has 3 suckers, while the right arm is Others like I. biserialis and I. more stocky and broad with 2 suckers at paradoxus inhabit sea grass and algae the base. areas. All observations on this genus DISCUSSION concerning behaviour, spawning, embryonic development and life cycle Systematics have only been made with wild-caught The relationships among species specimens in aquarium cultivation within this genus remain unknown. Jereb (Moynihan 1983; Jackson 1992). & Roper (2005) currently place eight Presently, relatively little is known about species within the genus: Idiosepius the biology and life cycles of Idiosepius. biserialis Voss, 1962, I. macrocheir For example, no data are available about Voss, 1962, I. minimus (D‘Orbigny, its postembryonic life, the onset of sexual 1845), I. notoides Berry, 1921, I. maturity or postembryonic behaviour. paradoxus (Ortmann, 1888), I. picteti Some authors assume a life cycle lasting (Joubin, 1894a), I. pygmaeus Steenstrup, 3 months (from egg development until 1881 and I. thailandicus Chotiyaputta death) (Tracey et al. 2003), but this value et al. 1991. needs verification. Moreover, the Morphologically the species can be geographical distributions of all identified by the arrangement of suckers Idiosepius taxa are still only marginally on the club (two or four rows) and the explored. This may be explained by their number of suckers on the ventral arms small size and habitat conditions: (hectocotyli) (Nesis 1982). observing specimens in their natural The species are mostly distributed habitat is still difficult. in the tropical Indo-Pacific, Japan, southern Australia including Tasmania, Habitat of Idiosepius in Indonesia and African waters, but individuals were Idiosepius biserialis was caught in also found in cooler Russian waters November 2007 at low tide with a dipnet

19 Byern & Marwoto

in two sea grass areas in the northern the genus Idiosepius and provide more area of Ekas-Bay, Lombok Island. These knowledge about their occurrence and observations agree well with other geographical distribution in Indonesian collections of I. biserialis in Africa, waters. Japan and Thailand, indicating that this species is specialised for sea grass areas ACKNOWLEDGMENTS (von Byern et al. 2005; von Byern & Klepal 2009). We still do not know We are very grateful to Didik whether the animals stay there at high Santoso, Lalu Japa & Karnan from tide or move elsewhere. In contrast, I. Mataram University, Biology Depart- pygmaeus could only be found in ment and furthermore to Mr. Mady mangrove forests and belts, such as those Marcuo and the Staff of Rinca Island for along the eastern part of Lombok Island helping collect the samples presented at Gili Sulat, Gili Lawang, Telong Elong here. Our thanks go in particular to Mrs. and Ekas-Bay, but are clearly absent in Toifl from the ASEA-UNINET of the the northern, western and south-western University of Vienna for promoting the shores of the Island. So far, no individuals research trip of the first author to and of this species were ever found in sea within Indonesia and Dr. Michael grass or algal habitats (Suwanmala et al. Stachowitsch from the University of 2006; von Byern & Klepal 2009). Vienna for critically reading the Moreover, individuals of the former manuscript. I. pygmaeus hebereri were collected in the eastern part of the Ekas-Bay in April REFERENCES 2004 (von Byern & Klepal 2007). During high tide, garbage, leaves and other plant Adam, W. 1986. La radula et les material were transported here by the mandibules de quelques espéces current. On some days this flotsam d´Idiosepius Steenstrup, 1881 covers almost the whole water surface. (Mollusca Cephalopoda Deca- Interestingly, the garbage in the water poda). Bull. de L´Institut Royal apparently does not affect I. pygmaeus: des Sci. Naturelles de Belgique even compact and strongly agitated 56: 149-154. waste had no influence on their Appellöf, A. 1898. Cephalopoden von behaviour and movement. Moreover, the Ternate. Abhandlungen der animals were also observed to mate Senckenbergischen Naturfors- within this flotsam and escape under the chenden Gesellschaft 24 (4): 570- waste when threatened. Their occurren- 637. ce between a flotsam of garbage indica- Berry, SS. 1921. Cephalopods of the tes the ability to adapt to new habitats. genera Sepiolidea, Sepiadarium, Additional genetic, ecological and and Idiosepius. The Philip. J. Sci. behavioral investigations are necessary 47 (1): 39-55. to provide a more complete picture of

20 Occurrence of Idiosepius (Mollusca: Cephalopoda)

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Memasukkan:Maret 2009 Diterima: Juli 2009