Opportunity to Explore the Speciation
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Odonatologica37(3): 235-245 September 1. 2008 The Fijian Nesobasis: a further examination of species diversity and abundance (Zygoptera: Coenagrionidae) H. Van Gossum¹’*,C.D. Beatty²,M. Tokota’a³ andT.N. Sherratt4 1 Evolutionary Ecology Group, University of Antwerp, Groenenborgerlaan 171, B-2020 Antwerp, Belgium 2 de de Grupo Ecologia Evolutiva, Departaraento Ecologia y BiologiaAnimal, Universidad de Vigo, EUET Forestal, Campus Universitario, ES-36005 Pontevedra, Galicia, Spain 3 International Conservation, South Pacific Program, 11 Ma’afu Street, Suva, Fiji 4 Department of Biology, Carleton University, 1125 Colonel By Drive, Ottawa, Ontario, K1S 5B6, Canada Received June 11, 2007 / Reviewed and AcceptedOctober 21, 2007 Recently, an overview of the diversity, abundance, distribution and morphological characteristics of spp. of the genus Nesobasis endemic to Fiji, was presented for , spp. occurring on the 2 largest islands of the archipelago: Viti Levu and Vanua Levu. Here, this knowledge is extended by providing more extensive diversity and abundance data for the island of Vanua Levu, as well as for 4 smaller islands in Fiji: Taveuni, Koro, Ovalau and Kadavu. Previous research indicated that the Nesobasis spp. inhabiting Viti Levu and Vanua Levu are unique, with these islands having no species in com- The data confirm this and also show that smaller islands in mon. new proposal prox- imity to these 2 larger islands usually contain a subset of the large island’s Nesobasis fauna. The island of Koro, however, is unusual in that, while its Nesobasis spp. are Vanua N. predominantlythose found on Levu, it also harbours rufostigma, a sp. oc- curring on Viti Levu. Further, N. recava is endemic to Kadavu and is not found on Viti Levu, the nearest large island. Species richness is higher on large than small is- lands while mean species abundances were consistently higher on large islands com- small islands. The of distribution and in this is pared to pattern speciation genus quite complex, and is the subject of ongoing research. INTRODUCTION The study of oceanic islands offers an excellent opportunity to explore the and of selection well the of process pattern as as nature speciation (DARWIN, * Corresponding author: [email protected] M. 236 H. van Gossum, C.D. Beatty, Tokota'a & T.N. Sherratt 1859; MacARTHUR & WILSON, 1967). Sadly, in many island groups, par- ticularly those in the Southwest Pacific, our knowledge of basic biodiversity is limited. Moreover, there is a fundamentallack of information on the behaviour, life history and ecology of endemic island fauna (WHITTAKER, 1998). In this information abundance context BEATTY et al. (2007) reported on the diversity, and basic morphology of species of the genus Nesobasis (Odonata: Zygoptera). Not is the Nesohasis endemic to it is also with only genus Fiji, species-rich, spe- cies distributed over several islands within Fiji (DONNELLY, 1990). Moreover, for of the within the males seldom observed at some species genus, are oviposi- tion sites (VAN GOSSUM et ah, 2007). This malerarity has not caused sex role reversal as previously speculated (DONNELLY, 1990, 1994) but research is on- going to explore what factors may explain male rarity in some species of Neso- basis but not in others (see also VAN GOSSUM et al., 2007). based research conducted in 2005 In our previous report, on (BEATTY et focussed Nesohasis observed Viti Levu and of Vanua al., 2007), we on on parts Levu. During our research in 2006 we were able to explore otherregions on Van- ua Levu and also to include informationon species diversity and abundance for four additionalsmaller islands: Kadavu, Koro, Ovalau and Taveuni. For these five islands we provide quantitative estimates of Nesohasis species diversity and abundance. Furthermore, as BEATTY et al. (2007) did for other species, we pro- characteristics identification videbasic morphological for field for one described for Uds We also species (N. recava) and a new species we encountered ( 3). make note of behaviour and any reproductive (tandem, copulation wheel, oviposition) observed for these species. In this paper we characterise the inter-island distribution of species of Nesoha- sis based primarily on our own observational data, but also on earlierpublished work (DONNELLY, 1990; BEATTY et al„ 2007). The islands of Taveuni and Koro are located closer to Vanua Levu (the second largest island in the archipel- ago) than Viti Levu (the largest island in the archipelago), while the islands of located Viti If Ovalau and Kadavu are nearer to Levu (Fig. 1). smaller islands share species with nearby larger islands, we expected the Nesohasis community on Taveuni and Koro to be a subset of species occurring on Vanua Levu, while Ovalau and Kadavu species were expected to occur also on Viti Levu. Many ecological factors can influence whether a disperser from the mainland establishes on an island. However, abundant species are more likely to produce higher absolute numbers of dispersers than rare species (MacARTHUR & WIL- SON, 1967; GASTON et al., 2006). Therefore, if dispersal played an important role in establishment that those determining success, one might expect species found on smaller islands would comprise a subset of the more abundant species on the large island. Species diversity and abundance in Nesohasis 237 METHODS the The damselflygenus Nesobasis is abundant and widespread in the Fijian islands. They are domi- nant Zygoptera in most stream habitats, particularly in mountain streams, but they are not found in rivers We used in combina- large or ponds (DONNELLY, 1990, 1994). topographic maps (1:50,000) tion with our prior experience with the genus (BEATTY et al., 2007) to select regionson islands that In D. were likely to contain appropriate Nesobasis habitat. addition, T.W. Donnelly and Polhemus communicated their earlier experiences with Nesobasis on the different islands. TheFiji Islands arelocated in the eastern Melanesian region of the South-western Pacific and have a warm, humid tropical maritime climate with mean monthly temperaturesfrom 23°C in Julyto 27°C in January (RYAN,2000). The islands aresubequatorialand exposed to heavier rainfall between No- vember and April, especially on the low islands and the leeward sides of the large islands (EVENHUIS & 6 BICKEL, 2005). Our study was conducted duringthe dryseasonin Fiji from August toOctober 4, 2006, atime of year similar to the timing of research summarised in BEATTY et al. (2007).Fiji consists 2 of several hundred islands and occupies a total area of approximately 650,000 km of which the land Levu area is less than 3 percent. There are two large islands in the archipelago (Fig. 1): Viti (10,388 2 2 km ) and Vanua Levu (5,535 km ) (see also EVENFIU1S & BICKEL, 2005). Viti Levu and Vanua with landforms volcanic Levu are rugged, including plugs, eroded calderas, deep gorges, and ravines carved by mountain streams, flat-bottomed valleys with extensive food plains, and mangrove dominat- ed deltas (EVENHUIS & BICKEL, 2005). Other islands in the archipelago areconsiderably smaller, manyhaving less pronounced topographyand lacking streams. However, several islands aresizable and predominantly volcanic in origin, having significant el- evations and most impor- tantly mountain streams that could hold species of Nesobasis. Of these smaller islands wesampled Kadavu 2 2 (408 km ), Koro (109 km ), 2 Ovalau (102 km ) and Tave- 2 uni (434 km ). Study sites that com- plied to our selection crite- ria (see further) for VAN- UA LEVU (see Fig. 1) in- clude: KR-01 (I6°30.79’S, 179°32.70’ E, alt. 265m), a large stream on the interi- or road above Korotesere, crossing approximately 1.2 km south of Seavaci Rd, we sampled upstream from the road; KR-02 (16°28.55’ S, 179°33.32’ E, alt. 170m), of six islands of for which data second stream above Ko- Fig. I. Map Fiji are presented, showing locations. For VANUA LEVU: - - rotesere, we sampled on sample (1) KR-01; (2) KR-02; — the downstream side of the (3) NagaciCreek; — (4) VunigautulailaiCreek; (5) WR-01. ForTA- road crossing; Nagaci Cre- VEUN1: (6)Somosomo; — (7) TV-02. For KORO: (8) Buretini Creek; - - ek (16°28.36’S, 179°35.32’ (9)K-01; (10) K-02. ForOVALAU: (11) Lovoni. For KADAVU: E, alt. 130m),small stream (12) Vunisea East. Site descriptions and latitude/longitudecoordinates are in the text. crossing road near small provided 238 H. van Gossum, C.D. Beatty, M Tokota'a & T.N. Sherratt settlement, we sampled downstream ofthe road; VunigautulailaiCreek (16°30.06’ S, 179°37.20’ E, alt. 110 m), stream at bridge on road above Vanuavou, we sampleddownstream of the road; WR-01 (16°41.12’ S, 178°55.29’E,alt. 335 m), river at high elevation in western Vanua Levu. For TAVEUNI (see Fig. 1) we sampled; Somosomo (16°46.40’ S, 179°58.22’ W, alt. 30 m), a large river at the south- ern edge of Somosomo village, we sampled upstream of the bridge; TV-02 (16°44.12’ S, I79°54.57’ 395 Somosomo We W, alt. m), small stream above in agricultural area. also sampled at Tavoro Falls, in Bouma National site that indicated be Nesobasis habitat Heritage Park, a was to very good (D. Polhemus, pers. com.). Despite searching an hour with two persons we were unable to locate any Nesobasis. For KORO (see Fig. l)we sampled Buretini Creek (I7°15.82’S, 179°22,08’E,alt. 150m), low astream with very flow, and predominantly exposed bedrock with boulders and numeroussmall pools, we sampled immediatelyabove a largewaterfall, working upstreamto the road crossing; K-01 (I7°15.82’ S, 179°22.08’ E, alt. 40 m), a stream along the road crossing the interior of the island in the north;K-02 (17° 19.82’ S, 179°25.92’ E, alt. 50 m), alarge stream on the east side of Koro, which crosses the main road south of the intersection with the interior road across the north side of the is- land.