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Diversity in Different Trophic Levels of the Plankton Biogeosciences Discuss., 5, 4897–4917, 2008 Biogeosciences www.biogeosciences-discuss.net/5/4897/2008/ Discussions BGD © Author(s) 2008. This work is distributed under 5, 4897–4917, 2008 the Creative Commons Attribution 3.0 License. Biogeosciences Discussions is the access reviewed discussion forum of Biogeosciences Diversity in different trophic levels of the Similar patterns of community plankton organization characterize distinct groups V. Raybaud et al. of different trophic levels in the plankton Title Page of the NW Mediterranean Sea Abstract Introduction V. Raybaud1,2, A. Tunin-Ley1,3, M. E. Ritchie4, and J. R. Dolan1,3 Conclusions References 1UPMC Univ Paris 6, UMR 7093, Laboratoire d’Oceanographie´ de Villefranche, Observatoire Tables Figures Oceanologique´ de Villefranche-sur-Mer, Station Zoologique, B.P. 28, 06230 Villefranche-Sur-Mer, France J I 2CNRS, UMR 7093, Laboratoire d’Oceanographie´ de Villefranche, Observatoire Oceanologique´ de Villefranche-sur-Mer, Station Zoologique, B.P. 28, 06230 J I Villefranche-Sur-Mer, France 3CNRS, UMR 7093, Laboratoire d’Oceanographie´ de Villefranche, Microbial Ecology and Back Close Biogeochemistry, Observatoire Oceanologique´ de Villefranche-sur-Mer, Station Zoologique, B.P. 28, 06230 Villefranche-Sur-Mer, France Full Screen / Esc 4Biology Department, Syracuse University, Syracuse, NY, USA Printer-friendly Version Received: 24 September 2008 – Accepted: 23 October 2008 – Published: 12 December 2008 Correspondence to: J. R. Dolan ([email protected]) Interactive Discussion Published by Copernicus Publications on behalf of the European Geosciences Union. 4897 Abstract BGD Planktonic populations were sampled over a 4 week period in the NW Mediterranean, at a site subject to little vertical advection during the Dynaproc 2 cruise in 2004. 5, 4897–4917, 2008 The characteristics of the phytoplankton, the tintinnid community and the zooplank- 5 ton have recently been described (Lasternas et al., 2008; Dolan et al., 2009; Ray- Diversity in different baud et al., 2008). Based on these studies we compared the characteristics of 3 trophic levels of the well-circumscribed groups of different trophic levels: Ceratium of the phytoplankton, plankton herbivorous tintinnids of the microzooplankton, and large (>500 µm) omnivorous and carnivorous copepods of the metazoan zooplankton. In all three groups, diversity as H0 V. Raybaud et al. 10 or species richness, was less variable than concentration of organisms. Plotting time against species accumulation, the curves approached plateau values for Ceratium spp, tintinnids and large copepods but only a small number of species were consistently Title Page present (core species) and these accounted for most of the populations. For Ceratium Abstract Introduction core species numbered 10, for tintinnids 11 species, and for large copepods, core 15 species numbered 4 during the day and 16 at night. Ceratium, tintinnids and large Conclusions References copepods showed some similar patterns of community structure in terms of species Tables Figures abundance distributions. Ceratium species were distributed in a log-normal pattern. Tintinnid species showed a log-series distribution. Large copepods communities were highly dominated with night samples showed much higher abundances and greater J I 20 species richness than day samples. However, species abundance distributions were J I similar between day and night and were mostly log-normal. The paradox of the plank- ton, describing phytoplankton communities as super-saturated with species, extends Back Close to the microzooplankton and zooplankton. Full Screen / Esc 1 Introduction Printer-friendly Version Interactive Discussion 25 Many groups of planktonic organisms are characterized by high species-richness. Hutchinson (1961) was the first to formally state that there appears to be an unrea- 4898 sonable number of phytoplankton species for an apparently homogenous environment – “the paradox of the plankton”. The paradox of a supersaturation of species has been BGD extended beyond phytoplankton to that of a general feature of aquatic systems (Roelke 5, 4897–4917, 2008 and Eldridge 2008). Over the years many solutions to the paradox, most applicable 5 only to phytoplankton, have been proposed but none have found general acceptance (Roy and Chattopadhyay, 2007). The answer may lie in examining mechanisms or Diversity in different phenomena which impact planktonic organisms in general. This possibility is difficult trophic levels of the to evaluate as a single distinct taxonomic or trophic group is nearly always examined plankton in isolation. There have been however, some studies comparing the large, very hetero- V. Raybaud et al. 10 geneous groups of “phytoplankton” and “zooplankton”. Interestingly, these few stud- ies which have examined “phytoplankton” and “zooplankton” agree in showing distinct differences. For example, different lake populations of phytoplankton and zooplank- Title Page ton appear to have different species abundance distributions (Walker and Cyr, 2007). However, the phytoplankton and zooplankton were not sampled in the same lakes. In Abstract Introduction 15 the marine plankton, phytoplankton diversity is a hump-shaped function of biomass along a large spatial gradient while zooplankton diversity appears to be a near-linear Conclusions References function of biomass (Irigoen et al., 2004). However, averaged over large time scales Tables Figures (decades) in a single large system – the Eastern Pacific Gyre, zooplankton and phy- toplankton show very similar species abundance distributions (McGowan and Walker, J I 20 1993). Phytoplankton and zooplankton diversity responds distinctly to disturbances, such as flushing, based on the results of experiments with lake plankton (Floder¨ and J I Sommer, 1999) and estuarine plankton (Buyakates and Roelke, 2005). Frustratingly Back Close few generalities about differences or similarities in plankton community structure have emerged, perhaps because both phytoplankton and zooplankton are heterogeneous Full Screen / Esc 25 groups, and phytoplankton are much more abundant and often more diverse than zoo- plankton. Printer-friendly Version Here we take a different approach to comparing phytoplankton and zooplankton, that of examining the diversity and community structure of circumscribed groups of Interactive Discussion species within the more general trophic classes of plankton. We profit from the si- 4899 multaneous in-depth studies of phytoplankton (Lasternas et al., 2008), zooplankton (Raybaud et al., 2008) and microzooplankton (Dolan et al., 2009) conducted during the BGD program “DYNAPROC 2”. We selected the large, generally omnivorous and carnivo- 5, 4897–4917, 2008 rous (>500 µm) copepods, Ceratium dinoflagellates of the phytoplankton, and herbivo- 5 rous tintinnid ciliates, to compare the characteristics of zooplankton, phytoplankton and microzooplankton. Each group is well-circumscribed, species-rich and one in which Diversity in different species identifications are straightforward. trophic levels of the Other than a planktonic existence, the assemblages have little in common. The plankton groups are trophically distinct: Ceratium species are dinoflagellate primary producers V. Raybaud et al. 10 containing chloroplasts. Tintinnids are ciliates, part of the herbivorous microzooplank- ton and feed mainly on small phytoplankton (5–25 µm in size). The large copepods (e.g. Neocalanus) are generally assumed to be omnivorous or carnivorous. The three Title Page sets of species vary in generation times with that of large copepods measured in weeks or months, Ceratium spp in days, and tintinnids in hours. The 3 groups also repre- Abstract Introduction 15 sent different degrees of phylogenetic cohesion: Ceratium are all obviously in a single genus; tintinnids represent a ciliate sub-order and “large copepods” groups species of Conclusions References distinct orders. The large copepod community differs as well from Ceratium and tintin- Tables Figures nids in that day and night communities in the surface layers are distinct with nighttime copepod communities constituted mainly of migrating taxa found in deeper waters dur- J I 20 ing the day. The three assemblages, whilst of distinct trophic levels, are unlikely to have direct impact on one another. The large copepods feed on Ceratium and tintinnid-size J I prey items but are not found in concentrations sufficient to affect Ceratium spp. or tintin- Back Close nids, given typical feeding rates for large copepods (e.g., Dagg et al., 2006). Ceratium spp., most of which exceed 100 µm, are too large to be ingested by most tintinnids. Full Screen / Esc 25 Here we compare the community characteristics of these sets organisms in terms of diversity, stability in terms of species identities and structure of community composition Printer-friendly Version by examining species abundance distributions and document the short-term variabili- ties. Interactive Discussion 4900 2 Material and method BGD 2.1 Sampling 5, 4897–4917, 2008 The Dynaproc 2 cruise, in September–October 2004, permitted repeated sampling over a 4 week period at a site in the NW Mediterranean Sea characterized by little ver- Diversity in different 5 tical advection. Detailed descriptions of the study site and sampling appear elsewhere; trophic levels of the here we will briefly review methods used to study copepods (Raybaud et al. 2008), plankton phytoplankton (Lasternas et al., 2008) and tintinnid ciliates (Dolan et al., 2008). The copepod data considered here are derived from
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