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The Systematics and Ecology of the Mangrove-Dwelling Littoraria Species (Gastropoda: Littorinidae) in the Indo-Pacific

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The Systematics and Ecology of the Mangrove-Dwelling Littoraria Species (Gastropoda: Littorinidae) in the Indo-Pacific ResearchOnline@JCU This file is part of the following reference: Reid, David Gordon (1984) The systematics and ecology of the mangrove-dwelling Littoraria species (Gastropoda: Littorinidae) in the Indo-Pacific. PhD thesis, James Cook University. Access to this file is available from: http://eprints.jcu.edu.au/24120/ The author has certified to JCU that they have made a reasonable effort to gain permission and acknowledge the owner of any third party copyright material included in this document. If you believe that this is not the case, please contact [email protected] and quote http://eprints.jcu.edu.au/24120/ THE SYSTEMATICS AND ECOLOGY OF THE MANGROVE-DWELLING LITTORARIA SPECIES (GASTROPODA: LITTORINIDAE) IN THE INDO-PACIFIC VOLUME II Thesis submitted by David Gordon REID MA (Cantab.) in May 1984 for the Degree of Doctor of Philosophy in the Department of Zoology at James Cook University of North Queensland 399 PART II: ECOLOGY CHAPTER 10: COCKLE BAY, THE PRINCIPAL STUDY AREA Magnetic Island is situated in Cleveland Bay, 7 km from the city of Townsville (Fig. 10.1). The island is approximately 5000 hectares in area and is mostly mountainous, reaching 506 m at its highest point. The geology of the island is comprised of acid plutonic rocks of late Palaeozoic age, with a narrow coastal plain of Quaternary deposits (Spenceley, 1982). The sheltered landward coast runs for 7 km in a north-westerly direction, separated from the mainland by a channel from 5 to 7 km in width. Along this coast an extensive mangrove forest has developed behind a broad mud flat with beds of sea grass, which extend to a fringing reef 1 to 1.5 km from the edge of the forest. At its widest point, the mangrove forest extends for 500 m, and adjoins a MeVaLeuca swamp or Eucalyptus woodland at the level of the highest spring tides. The floristic zones of seaward Avicennta fringe, RhtzophOra forest, Brugutera forest and Certops thickets are all represented (Section 11.2; Macnae, 1968), but the Certops zone is in some parts interrupted by a bare salt flat (Fig. 11.2). The Vegetation of the island coast has been described in detail by Macnae (1967) and Spenceley (1982). Climatic and tidal data for the Townsville area are summarized by Spenceley (1982). The rainfall is highly seasonal, 76% of the average annual total falling between January and April. An annual rainfall of 1825 mm (recorded at Picnic Bay, 1 km from the main study site) has permitted a relatively luxuriant development of mangrove vegetation on the landward side of Magnetic Island. In contrast, the mainland coast of Cleveland Bay has a lower rainfall (1163 mm at Townsville, Spenceley, 1982) and mangroves are present Fig. 10.1 Map of Magnetic Island and Cleveland Bay, Queensland. Dotted line indicates extreme low water of spring, tides. Fig. 10.2 Map of the study area at Cockle Bay, Magnetic Island, Queensland. a, line of transect represented in Fig. 11.1; b, 15 by 15 m study area in Rhtzophora grove; c and d, study areas (defoliated and normal Avtcennta bushes respectively) for investigation of frequency-dependent selection on colour forms of Ltttorarta ftLosa (Section 15.3); e, limits of experimental zone; f, sampling zone; solid asterisks, sites of five Avtcennta trees for mark and recapture study of L. phtttpptana; open asterisks, sites of two Avtcennta trees for mark and recapture study of L. ftLasa; light stipple, sclerophyll woodland; heavy stipple, mangrove forest. 146°50'E 190 10'S - c" Picnic Bay Cockle Bay CLEVELAND BAY TOWNSVILLE O 5 km • 402 mainly as narrow fringes to extensive bare salt pans (Macnae, 1967). It was for this reason that Magnetic Island was chosen as the main study area. Furthermore, the land behind the mangrove coast is virtually uninhabited, and the area should be less affected by domestic, industrial and agricultural pollution than the mainland coast. Several Ltttorarta species are scarce or absent on muddy continental coasts, and more common in narrow mangrove fringes and where the coastal water is less turbid (Section 11.4.5). Thus, while only three species were to be found in the broad and muddy forest midway along the mangrove coast of Magnetic Island (as listed, Fig. 11.2), five species were common at Cockle Bay at the southern end of the belt of forest (Fig. 11.1). Cockle Bay was therefore selected as the study site, (19° 10.5'S 146° 49.6'E; Fig. 10.2). The structure of the forest at Cockle Bay is somewhat atypical, by virtue of a long sand bar which has developed parallel to the coast, within what was once a belt of Rhtzophora forest. As a result, the typical zonation pattern of the forest has been interrupted. At the present time there are several isolated groves of Rhtzophora styLosa trees, up to 8 m in height, in front of the dune, and behind them an open area of muddy sand with a few isolated trees of Avtcennta eucaLypttfoLta and increasing numbers of colonizing Avtcennta bushes and saplings (Figs 10.3, 10.4). A transect through this area is represented in Fig. 11.1. The study area was divided into experimental, sampling and undisturbed zones (Fig. 10.2). Within the experimental zone a number of trees were selected in three areas, on the basis of the abundance of four species of Ltttorarta which were to be studied in detail. The first area comprised all the 64 Rhtzophora trees in the space of 15 m by 15 m within a Rhtzophora grove, where L. scabra, L. tntermedta and L. pht7Apptana were common. Secondly, two isolated trees of Avtcennta between the Rhtzophora grove and sand bar, were chosen because of the abundance of L. ftLosa. Thirdly, five larger Fig. 10.3 The exclusion cages at Cockle Bay, Magnetic Island, Queensland, See Section 14.2.2 for details. The large Avtcennta tree behind the foremost cage is one of the two on which the mark and recapture study of •ittorarta ftLosa was carried out. Fig. 10.4 Aerial view of study area at Cockle Bay, Magnetic Island, Queensland. Road to Picnic Bay in background (see map, Fig. 10.2). 405 Avtcennta trees on the edge of the sand bar were selected for study, on each of which L. phtLipptana was common. Details of the dimensions of trees are given in Table 10.1. All trees were identified individually by numbers, and the ground level beneath each was estimated (to the nearest 0.1 m) in relation to tidal datum, by comparison of water depth with predicted tidal heights at recorded times on three occasions. Within each of these study areas all the snails were individually numbered, for the purposes of recording their location on the trees, their growth and, by employing a mark and recapture technique, to investigate population dynamics. Details of the methods used are given in the appropriate later chapters. Exclusion experiments, to measure loss due to predation and other causes, were carried out on small, isolated Avtcennta trees in the area between the Rhtzophora grove and the sand bar (Fig. 10.3). An experiment to investigate frequency-dependent selection was performed in an area with numerous isolated and equally sized Avtcennta bushes (Fig. 10.2). Specimens for the examination of reproductive condition were collected from the sampling area reserved for the purpose. Table 10.1 Dimensions of trees in three main study areas at Cockle Bay, Magnetic Island, Queensland. Ltttorarta sp. Tree sp. No. of Height Lowest Canopy Diameter studied trees (m) foliage width at breast (m) (m) ht. (cm) L. scabra, Rhtzoptiora 64 4-8 1-3.5 1-3 5-18 L. tntermodia, styLosa L. phtLtpptana L. ftLosa Avtcennta 2 3,4 0.6,0.1 3 5,10 eucaLypttfoLta L. phiLtppiana Avicennta 5 4-5 0.2-1.4 3-5 10-17 eucaLypttfoLta 407 CHAPTER 11: HABITAT AND ZONATION 11.1 Introduction The association of many members of the genus Ltttorarta with mangrove, salt marsh and drift wood habitats represents amajor adaptive shift away from the ancestral habitat of intertidal rocks. That this condition is indeed derived is indicated by the fact that Ltttorarta species typical of rocky substrates occur only in the paraphyletic nominate subgenus (Section 8.1.2 and Fig. 8.2). Outgroup comparison with the sister group Nodtlittortna leads to the same conclusion, for all but one of the many species of NodtZtttortna occur only on rocky shores (pers. obs. and museum collections). In littorinid genera other than Ltttorarta, only a few species can be found on vegetation. These include a single species of NodtLittortna (a rare, undescribed species from South East Asia), Bembtctum meLanostoma (in eastern Australia), a species of PeasteLLa (in northern Australia) and several members of the genus Ltttortna which are sometimes found on salt marsh vegetation in the northern Atlantic (Raffaelli, 1978b). At most mangrove localities Ltttorarta species are the only abundant arboreal gastropods. On the trees at the seaward edge of the forest, gastropods typical of rocky shores can sometimes be found, including species of PateLLotda, Monodonta, CLypeomorus, Planaxts, MoruLa and Thats, usually at low densities and at tidal levels below those occupied by Ltttorarta. Within the forest several species of Nertta are found, while towards the landward edge of the forest Certthtdea and various ellobiid pulmonates climb the mangrove trees, descending to feed at low tide (see Macnae, 1968; Berry, 1972; Frith et aL., 1976, for general accounts of the molluscan fauna of Indo-Pacific mangroves). At none of the localities visited during the present study did these other arboreal gastropods occur 408 in large numbers at the same levels as Ltttorarta species, and their respective patterns of distribution did not suggest any evidence of interspecific competitive effects.
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