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Seamounts, Hotspots for High Speciation Rates in Bentho-Pelagic Fishes? a Case Study on Macroramphosus Spp

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Seamounts, Hotspots for High Speciation Rates in Bentho-Pelagic Fishes? a Case Study on Macroramphosus Spp Matthiessen et al., Macroramphosus CM 2002/M: 07 1 Ices CM 2002 / Oceanography and Ecology of Seamounts - Indications of Unique Ecosystems (Session M) Not to be cited without prior reference to the author. Seamounts, hotspots for high speciation rates in bentho-pelagic fishes? A case study on Macroramphosus spp. (Syngnathidae) from Great Meteor Seamount BIRTE MATTHIESSEN, HEINO O. FOCK, HEIN VON WESTERNHAGEN Alfred-Wegener-Institut für Polar- und Meeresforschung, Am Handelshafen 12, D-27570 Bremerhaven, Germany, Phone: 49-0471-4831 1384, [email protected] Snipefishes of the genus Macroramphosus (Syngnathidae) are the most abundant demersal fish species on the Great Meteor Seamount (GMR, subtropical NE Atlantic, 30° N, 28,5° W). High morphological variability within the genus at GMR questions the presence of only one species of Macroramphosus. Thus, 202 individuals of Macroramphosus spp. were investigated with respect to diet composition and morphology. A deep-bodied benthos feeding type (b-types) whose diet consisted of foraminiferans, pteropods, decapods and polychaetes, could be distinguished from a slender planktivorous type (p-types, n=140) mainly feeding on ostracods, copepods, pteropods and foraminiferans. Few specimens showed no specialized feeding mode (p/b-types). Both feeding types showed significantly morphologically differences after applying bi- and multivariate statistical analysis considering the variables body depth, length of second dorsal spine, diameter of orbit and standard length. Based on a randomisation test, the analysis of dietary overlaps revealed significant differences between feeding types. This result reinforces the hypothesis of two competing species of snipefishes at GMR. Notwithstanding similar observations on snipefishes from shelf habitats from the Pacific, our results suggest that isolated seamounts may represent hotspots of speciation where special selection pressure favours the evolution of two ecological divergent species within the genus Macroramphosus. Keywords: Macroramphosus, Great Meteor Seamount, habitat partitioning, sympatric species, null-model, randomisation, competition Matthiessen et al., Macroramphosus CM 2002/M: 07 2 Introduction In all oceans, snipefishes of the genus Macroramphosus Lacepède 1803 (Centriscidae, Macroramphosinae)1 abundantly inhabit shelf regions and seamounts, principally between 20 and 40° of latitude in both hemispheres (e.g. Parin et al., 1997; Wilson and Kaufmann, 1987). Two vernacular names have been established for Macroramphosus spp. to account for a world-wide observed variability, i.e. longspine snipefish and slender snipefish. At present, the world-wide record of thirteen species can be traced back to these two different types of species, i.e. the longspine snipefish M. scolopax (L. 1758, originally described as Balistes scolopax) and the slender snipefish M. gracilis (Lowe 1839, originally described as Centriscus gracilis) (Eschmeyer, 1998). Though present in all oceans, the taxonomic status for M. gracilis is still unclear, since it is both treated as a genuine species and as a juvenile form of M. scolopax (see Froese and Pauly, 2001; Mohr 1937). A review of the status of Macroramposus in the NE Atlantic by Ehrich and John (1973) initially distinguished two distinct species in the North Atlantic. However, the findings of intermediate specimens then forced Ehrich (1976) to revise the status of M. gracilis so that it became a juvenile stage of M. scolopax. Consequently, Ehrich (1976) subsumed all his specimens of the genus under M. scolopax. Clarke (1984) investigated Macroramphosus spp. from near the edge of the continental shelf of SE Australia. Due to two distinct dietary types which also differed significantly in morphological characters, he postulated the existence of two separate species of Macroramphosus in SE Australia. In turn, he also discussed whether this could resemble the variability in other parts of the world's oceans as well. A further analysis of recent material from GMR applying methodologies of both Ehrich (1976) and Clarke (1984) revealed a strong dependence of morphology on feeding type and thus supported Matthiessen et al., Macroramphosus CM 2002/M: 07 3 Clarke's (1984) hypothesis, indicating that two sympatric species of Macroramphosus were also present in the NE-Atlantic (Matthiessen et al., submitted). Selection pressure by means of competition promotes divergence in adaptive radiation (Schluter, 1994). Thus, new exploitative capabilities are attained (Schluter, 1993). The amount of overlap in resource utilization is assumed to be proportional to the intensity of competition between two species (Schoener, 1974a). In turn, morphologically similar species are likely to be stronger competitors than morphologically different species (Juliano and Lawton, 1990; Ricklefs et al., 1981; Warren and Lawton, 1987). Sympatric competing members of a guild often vary in decisive morphological features by a ratio known as Hutchinson's rule in order to partition diet and habitat (Schoener, 1974a). Hence, the exploitation of new resources and the associated interspecific differences in morphology are both signatures of adaptive radiation (Futuyama, 1990). This rationale paves the way for a synoptic investigation strategy considering both the exploitation of resources, i.e. diet, as well as conjoint differences in morphology and diet composition. Significant results of conjoint morpho-dietary differences were already analysed and are summarised by Matthiessen et al. (submitted). Further, competition theory warrants significant differences in diet utilisation between the observed morpho-dietary types in order to evidence sufficient specific separation. This is investigated in this paper by means of randomisation procedure. To analyse the observed pattern of morphologically distinct diet-types (benthic and planktonic feeders) with regard to resource partitioning and limiting competition it is useful to compare the observed data with an assumed pattern-free randomised pseudo- community. The central prediction of ecological and evolutionary models of displacement is that species should overlap less in resource use than in the absence of competition (Schoener, 1974a; Schoener, 1974b). There is wide support for the theory 1 Nomenclature follows Eschmeyer (1998). Matthiessen et al., Macroramphosus CM 2002/M: 07 4 that similar species reduce interspecific competition by subdividing their use of available resources (Pianka, 1974; Schoener, 1974a; Winemiller and Pianka, 1990). By making use of morphological and dietary analysis within the genus Macroramphosus we are trying to elucidate the existence of two sympatric species of snipefishes on Great Meteor Seamount. Materials & Methods Sampling For the study Macroramphosus from the Great Meteor Seamount (subtropical NE Atlantic, 30°N 28.5°W) obtained during the M42/3-cruise of the RV Meteor in September 1998 (Pfannkuche et al., 2000) was analysed. Sampling was conducted with a GOV-bottom trawl with 450 meshes opening width and 32 m wing span (Uiblein, 1999); detailed description in Fock et al. (2002b). Stretched codend mesh size was about 15 mm. Sampling depth varied between 295 to 349 m. On Great Meteor Seamount Macroramphosus spp. and Capros aper were the numerically dominant fish species (Uiblein, 1999). For Macroramphosus spp. a total of 82700 fish were caught. For the extensive analyses of morphology and diet, 202 fishes were randomly drawn from deep- frozen sub-samples with the intension to cover a diurnal cycle (Table 1). Morphometrics In order to generate data which were comparable to earlier investigations (Clarke, 1984; Ehrich, 1976) the same morphometric characters were measured (Fig. 1). Matthiessen et al., Macroramphosus CM 2002/M: 07 5 Analysis of diet and dietary overlap Due to lack of a well defined stomach, the whole alimentary tract was examined for dietary analysis. Following Clarke (1984) the alimentary tract was separated into an anterior, mid and posterior section and the rectum. Prey items were identified to major taxa by means of a dissection microscope. Items with fleshy tissue were counted as dietary items, items without any flesh were counted as remains, e.g. single echinoderm spines, single polychaete setae, calcareous deposits of unknown origin. With regard to the main diet, specimens were classified as either plankton feeders (p-type), benthic feeders (b-type) or mixed feeders (p/b-type) after Clarke (1984). Length-frequency distributions were applied to check whether comparable size ranges of the fishes were obtained within different categories compared to the total population on the Great Meteor Seamount. Distributions were based on total length measurements, since total length was measured onboard during the 1998-cruise. For each prey item an `Index of Relative Importance` (IRI) was calculated after Hyslop (1980), based on numbers (N), frequencies (F) and weight (W): 100 Ai IRIi [%] = n ∑ Ai i with Ai = % Fi + % N i + %Wi , and Fi (%) = Percentage of guts with food item i of the total number of analysed guts Ni (%) = Percentage of food item i of the total number of food items Wi (%) = Percentage of ashfree dryweight (AFDW) of food item i of the total AFDW n = Number of different food items Matthiessen et al., Macroramphosus CM 2002/M: 07 6 Empty alimentary tracts were included in the calculation. Statistical analysis of dietary overlap The analysis of dietary overlap (Gotelli and Graves, 1996; Lawlor, 1980; Winemiller and Pianka, 1990) was carried
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