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(Penaeus Indicus H. Milne Edwards, 1837) and Juvenile White Banana

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(Penaeus Indicus H. Milne Edwards, 1837) and Juvenile White Banana Journal of Experimental Marine Biology and Ecology 309 (2004) 79–108 www.elsevier.com/locate/jembe Allopatric distribution of juvenile red-legged banana prawns (Penaeus indicus H. Milne Edwards, 1837) and juvenile white banana prawns (Penaeus merguiensis De Man, 1888), and inferred extensive migration, in the Joseph Bonaparte Gulf, northwest Australia R.A. Kenyona,*, N.R. Loneragana, F.J. Mansona, D.J. Vancea, W.N. Venablesb a CSIRO Marine Research, P.O. Box 120, Cleveland QLD 4163, Australia b CSIRO Mathematical and Information Sciences, P.O. Box 120, Cleveland QLD 4163, Australia Received 4 August 2003; received in revised form 9 March 2004; accepted 11 March 2004 Abstract During October to December 1997, we trawled estuarine habitats in the Joseph Bonaparte Gulf (JBG) to determine the distribution of juvenile red-legged banana prawns, Penaeus indicus (H. Milne Edwards, 1837) and white banana prawns, Penaeus merguiensis (de Man, 1888). We made 229 beam-trawls at 185 sites, mostly over a 100-m path (3-min duration). A Global Positioning System (GPS) receiver was used to verify our location. During October to December 1998, we intensively resampled three of the rivers that were sampled in 1997 to confirm the gulf-wide distribution of P. indicus and P. merguiensis and to investigate the microhabitat use of P. indicus. We chose previously sampled and new sites in Forsyth Creek (eastern JBG), the Lyne River (Cambridge Gulf), and the Berkeley River (western JBG). We made 249 trawls at 21 sites, mostly over 100 m. Juvenile banana prawns were abundant in eastern JBG, Cambridge Gulf and western JBG. They were not abundant in southern JBG, although fewer trawls were made there, due to its inaccessibility. In eastern JBG and Cambridge Gulf, over 96% and 73% (respectively) of juvenile banana prawns were P. indicus and they were more abundant there than in the western JBG. Conversely, in the western JBG over 93% of the juvenile banana prawns were P. merguiensis and they were more abundant than in the eastern JBG and Cambridge Gulf. The Lyne River in the northwestern * Corresponding author. Tel.: +61-7-3826-7274; fax: +61-7-3826-7222. E-mail address: [email protected] (R.A. Kenyon). 0022-0981/$ - see front matter D 2004 Elsevier B.V. All rights reserved. doi:10.1016/j.jembe.2004.03.012 80 R.A. Kenyon et al. / J. Exp. Mar. Biol. Ecol. 309 (2004) 79–108 Cambridge Gulf seems to be the transition zone; both P. indicus and P. merguiensis are equally abundant. P. indicus are most abundant on the mangrove-lined muddy banks of waterways within mangrove forests, similar habitats to P. merguiensis. Within these habitats, they were most abundant in gutters and small creeks, rather than rivers and large creeks. Few P. indicus or P. merguiensis were caught in 100 m2 trawls undertaken midriver (on the channel bottom and on emergent banks), although these habitats may be only 100 m from the mangrove-lined habitats. In all creek and river habitats, both species are most catchable at low tide (irrespective of daylight or darkness) when they move out of the mangrove forests and accumulate in the remnant water bodies. The offshore fishery for P. indicus is in northwestern JBG in waters 50–80 m deep, about 300 and 200 km, respectively, from where juveniles are abundant in their extensive inshore habitats in east JBG and in Cambridge Gulf, demonstrating a geographical separation of the juvenile and adult phases. Postlarval P. indicus, spawned offshore, must use tides and currents to travel south and east to reach nursery habitats. Emigrant subadults must migrate north and west, across relatively shallow inshore sand substrates (30–40 m deep) to reach their offshore habitats. D 2004 Elsevier B.V. All rights reserved. Keywords: Hydrodynamic processes; Mangrove habitat; Microhabitat; Penaeus indicus; Penaeus merguiensis 1. Introduction Throughout the coastal regions bounding the Indian and western Pacific Oceans, the banana prawns (shrimp) Penaeus indicus and Penaeus merguiensis exist in sympatric inshore and offshore habitats (Devi, 1988; Rao et al., 1993; Barus and Mahiswara, 1994; Sambandam, 1994; Mohan et al., 1995; Primavera, 1996). Both species have a similar life cycle; the adults spawn offshore, pelagic larvae migrate inshore, juveniles spend several months in mangrove-lined estuaries and then they migrate offshore (Dall et al., 1990). They support significant coastal and estuarine fisheries adjacent to estuarine mangrove nursery areas (Rao et al., 1993; Mohan and Siddeek, 1995, 1996). Sometimes, juvenile P. indicus and P. merguiensis occupy similar microhabitats among soft-sediment mangrove communities (Mohan et al., 1995). In Australia, the population biology of P. merguiensis has been studied intensively for over 20 years (e.g., Crocos and Kerr, 1983; Vance et al., 1985; Staples and Vance, 1986; Vance et al., 1998). The juveniles are abundant in mangrove-lined estuaries. Their abundance is greatest in the upper reaches of small creeks (Vance et al., 1998), especially among mangrove forests with many small streams probably where the physical complexity of the habitat is greatest (Webb and Kneib, 2002). In contrast, the habitat use and distribution of juvenile P. indicus in Australia is relatively unknown (Somers, 1994). In other countries within their range, juvenile P. indicus occur on muddy sediments in creeks and rivers among estuarine mangrove forests (Subramanian, 1985; Mohan and Siddeek, 1996), where they are tolerant of euryhaline conditions (Subramanian, 1985; Kumlu and Jones, 1995). Often, subadult and adult P. indicus are found over a range of habitats, from coastal bays to inshore and offshore waters where they are commercially exploited at different stages of their life R.A. Kenyon et al. / J. Exp. Mar. Biol. Ecol. 309 (2004) 79–108 81 history (George et al., 1963; Rao et al., 1993; Mohan and Siddeek, 1995). Historically, in India, P. indicus were fished in 10–40 m depths, while more recently larger vessels allow exploitation in depths up to 100 m (George et al., 1963; Suseelan and Pillai, 1993).In Madagascar, the adults are found close to shore in 15–25 m water depth (Brinca and Mascarenhas, 1985). In Australia, only the offshore adults are fished in waters about 50– 80 m deep. Compared to other tropical penaeids in Australia, P. indicus are found in a very restricted geographic range—the Kimberley and Arnhem Land coasts, and the east coast of Cape York (Grey et al., 1983). They do not extend beyond 20jS on the west or the east Australian coasts, and within this range they are often uncommon. Their limited extent in northern Australia reflects their existence on the southeastern limits of their global zoogeographic distribution. The Timor, Arafura and Coral Seas are the eastern extent of the range of the species, which flourishes in the coastal seas along the western and northern Indian Ocean (Dall et al., 1990). As well, their distribution in northern Australia may be reduced by competition with P. merguiensis, a similar species which dominates mangrove habitats in the western Pacific Ocean and flourishes in the eastern and northern Indian Ocean (Dall et al., 1990). P. indicus are fished in the western Joseph Bonaparte Gulf (JBG) (Fig. 1) and north of Melville Island (11.5jS, 131jE)/Coburg Peninsula, the only Australian locations where they are commercially abundant (Somers, 1994). In contrast to P. indicus, the white banana prawn, P. merguiensis, is found over a wide area of northern Australia, from south of Moreton Bay (27jS, 153.25jE) on the Fig. 1. Study sites trawled for postlarval and juvenile banana prawns during three surveys undertaken in the JBG during October and December 1997. 82 R.A. Kenyon et al. / J. Exp. Mar. Biol. Ecol. 309 (2004) 79–108 east coast, to Shark Bay (25.5jS, 113.5jE) on the west coast (Grey et al., 1983).In Australia, P. merguiensis are common and fished commercially throughout their range (Somers, 1994) in locations offshore to their estuarine mangrove nursery habitats (Staples et al., 1985). In the JBG, the geographic distributions of the dominant P. indicus fishery and the P. merguiensis fishery do not overlap (Somers, 1994). P. merguiensis are fished from the northwestern extremity of the JBG, westward to Napier Broome Bay (14jS, 126.5jE). The location of the P. indicus fishery in the north western JBG and extensive areas of estuarine habitats (potential nursery habitats) in the southeast JBG, distant from the fishery, raises questions as to the coastal distribution of the juvenile P. indicus that recruit to the fishery. Usually, juvenile banana prawn habitats are adjacent to the fishery, in inshore estuarine habitats (Staples et al., 1985). If the juveniles of both banana prawn species were distributed evenly throughout coastal JBG, why were both species not common in the fishery catch? Was only one species common in the nursery habitats of the JBG? Until recently, juvenile P. indicus were very difficult to separate from P. merguiensis using morphological characters making their study difficult in areas where their distributions overlapped (Pendrey et al., 1999). In this study, our aim was to investigate the broad geographic distribution and abundance of juvenile P. in dic us and P. merguiensis in the coastal and estuarine habitats along 500 km of the JBG. We intended to determine if both species were common throughout all regions, or if few P. merguiensis were found in the JBG. In addition, we studied the habitat use of both species in three regions of the JBG to test for difference in microhabitat use between P. indicus and P. merguiensis. To complete these aims, features of their mangrove habitats were quantified and used to explain the distribution of the juveniles as found. As well, the effects of other environmental factors on the distribution patterns were investigated using a quasilikelihood generalised linear model.
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