Latitudinal biodiversity patterns of meiofauna from sandy littoral beaches
Contents
Introduction
This article is about latitudinal biodiversity patterns. The latitudinal biodiversity pattern of the meiofauna of sandy littoral beaches is discussed as a special case.
Latitudinal distribution of marine biodiversity
Large-scale patterns of marine biodiversity have been subject of many discussions (e.g.[1] [2] [3] [4] [5] [6] [7] [8] [9] [10]). These attempts to develop a general picture of diversity in the sea are hampered by the small number of key studies, the varied sampling protocols applied, the different diversity indices and the varying levels of taxonomic resolution [11]. A general trend of species impoverishment towards the poles was reported for some taxa (e.g., corals, gastropods), but this does not hold for others (e.g., amphipods and decapods crustaceans) [12] [13] [14]. In addition, it seems probable that there is a cline in increasing diversity from the arctic to the tropics, but the cline from the Antarctic to the tropics is far less well-established [6] since the Antarctic has high diversity for many taxa [15] [16]. Broad latitudinal gradients in species richness are illustrated for open-ocean pelagic and deep-sea taxa, but some debate continues to surround evidence for shallow-water systems, particularly for non-calcareous taxa [17]. Gray (1997)[6] stated that marine biodiversity is higher in benthic (bottom-related) rather than in pelagic (in the water column) systems, and on coasts rather than in the open ocean, since there is a greater range of habitats near the coasts. A good comparison of multispecies macrofaunal assemblages inhabiting the same type of habitat (sublittoral, fine sediment bottom) showed little if any difference among tropical, temperate and arctic sites in terms of diversity[18]. There was some dispute on how far the observed latitudinal patterns are size-dependant and small bodied taxa (Protoza and meiofauna) tend to be more ubiquitous and their richness is less latitude dependant compared to large organisms[19][20][10].
In coastal environments the interactions between coastal morphology, land-ocean exchanges, meteorological and tidal conditions, create a highly complex and finely scaled network of environmental boundaries. These boundary conditions explain why coastal waters have both higher species richness and a richer ecosystem than their oceanic counterparts [3]. Sandy shores are among the most ‘simple’ systems in terms of habitat complexity in comparison to other coastal ecosystems as, for example, rocky shores, algae and seagrass beds. Biodiversity and biomass of interstitial organisms are rather low. However, recent findings have shown that marine sands transfer energy very effectively, and that chemical and biological reactions take place faster there than in fine-grained sediments[21].
Latitudinal distribution of meiofauna
The maximum meiofauna densities reported in the study of Kotwicki et. al (2005)[22] ranged between 15 and 4312 individuals per [math]10 cm^2[/math]. The reported densities rank among the meiofauna densities in sandy beaches reported in available literature. In general, high meiofauna density can be found in intertidal muddy estuarine habitats, while much lower values are recorded in the deep sea [23]. In fine sediments such as organic rich muds, meiofauna densities of 104 individuals 10 cm sq and more are common [24]. The available meiofauna data showed a large within-site (within-region) variation in the temperate zone while there was very little variation within the meiofauna densities of the antarctic and arctic zones[22]. These patterns demonstrate that attempts to project global biodiversity from the results of regionally based studies must include the significant variation in diversity among sampling sites. The low salinity effect on meiofauna occurrence was not clear – two brackish water locations have the same range of meiofauna density as full marine sites[22]. As was reported in cited literature, the lower salinity was not associated with decrease of meiofauna[25].
In terms of taxonomical composition, the meiofauna taxa that were encountered during study of Kotwicki et. al (2005)[22] are similar to those of muddy sediments. There were no meiofauna taxa found that were restricted to shallow permeable sediments only. The percentage composition, on the other hand, differed significantly between the different study sites along the latitudinal gradient. In general, nematodes dominate benthic meiofauna communities comprising more than half of the total meiofauna abundance. This was indeed the case for most sampling sites except for both polar regions (arctic and antarctic), where turbellarians were the dominant meiofauna group.
Some small macrofaunal crustacean species (Cumacea, Amphipoda, Mysidacea) that can occasionally be found in meiofauna samples, were absent in the littoral zone of polar waters[22]. Small size in macrofauna is often associated with a fast development, r-strategy and warm environmental conditions that permit fast egg incubation and growth[26]. That strategy is unlikely to be fruitful in cold regions. These results support the hypothesis that warm regions support fast growing, smaller and more abundant organisms, and cold regions are dominated by larger and less abundant meiofauna. Macrofauna taxa may contribute to the meiofauna size class only in the tropics. A lower number of taxa was collected in both polar sites and, this in combination with slightly higher diversity in the temperate and tropic zones, supported the general pattern of diversity increase towards lower latitude. When only ‘meiofauna sensu stricto’ (i.e., without the small macrofaunal organisms) were taken into account, no clear latitudinal change could be found.
Meiofauna biodiversity of sandy shores
However, data at species level can give a more detailed and perhaps different outcome. Average number of nematode species from sandy littoral sites ranges between 50 and 60 species in warm temperate localities (Italy), cold temperate (Baltic) and slightly less are reported from arctic Svalbard (Gheskiere, Ghent University, personal communication). The results of the classification illustrated the clear difference between the polar sampling sites on the one hand and the more temperate beaches on the other hand. Archambault and Bourget (1996)[27] showed that large-scale heterogeneity explains a larger proportion of the variance in macrofauna species richness than substratum heterogeneity on a more local scale. In this context, it is fairly reasonable to refer to the important human pressure on temperate beaches. Recent studies (e.g.[28] [29]) focused on the effects of recreational pressure (trampling, beach cleaning and nourishment). Weslawski et al. (2000)[28] suggested that a highly diverse meiofauna and diatom assemblage in undisturbed beaches may act as an effective biological filter for some types of pollutant, while less diverse, but more abundant biota in disturbed areas are more effective in processing organic matter (self-cleaning of the beach). Largely as a result of conflicting uses of coastal habitats, losses of marine diversity are highest in coastal areas. The best way to conserve marine diversity is to conserve habitat and landscape diversity in the coastal area. Marine protected areas are only a part of the conservation strategy needed. A framework for coastal conservation should include integrated coastal area management, where one of the primary goals is sustainable use of coastal biodiversity[28].
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