Meiobenthic diversity in space and time: The case of harpacticoid copepods in two
Mediterranean microtidal sandy beaches
Katerina Sevastou
a,
⁎, Nikolaos Lampadariou
b
, Anastasios Eleftheriou
a, c
a
Department of Biology, University of Crete, P.O. Box 2208, 71409 Heraklion, Crete, Greece
b
Hellenic Centre for Marine Research, Institute of Oceanography, P.O. Box 2214, 71003 Heraklion, Crete, Greece
c
Hellenic Centre for Marine Research, Institute of Marine Biology and Genetics, P.O. Box 2214, 71003 Heraklion, Crete, Greece
abstract article info
Article history:
Received 20 January 2011
Received in revised form 24 June 2011
Accepted 28 June 2011
Available online 12 July 2011
Keywords:
Meiobenthic Copepods
Alpha Diversity
Beta Diversity
Turnover Diversity
Sandy Beaches
Eastern Mediterranean
Meiobenthic data from two microtidal sandy beaches of the eastern Mediterranean (Crete, Greece) were used
to investigate patterns of both alpha and beta diversity in space and time. Copepod assemblages and
environmental variables related to sediment characteristics, morphodynamics and food were studied over a
year at four distinct habitats at each beach; the retention, resurgence and saturation zones of Salvat's
intertidal scheme (midlittoral zone), and the surf zone of the sublittoral. Αlpha diversity analysis indicated
similar species richness at both beaches when the whole 13-month data set was considered but was higher at
the sheltered site when each sampling period was examined separately. Both beaches supported higher
diversity in the sublittoral zone. Species richness increased seawards at the midlittoral zone of the sheltered
site whereas, no pattern was evident at the exposed site, where the intense hydrodynamic conditions
homogenized the sediments. Beta diversity increased markedly towards the sublittoral, indicating greater
differences in alpha diversity between the sublittoral and the midlittoral zone. Species turnover was more
variable at the exposed beach and at the most landward stations, where environmental conditions change
often between extremes. A proportion of the variation in alpha diversity was explained by food availability at
both beaches and additionally by grain size at the sheltered site. However, no environmental variable
explained beta diversity patterns. Although the results of our study support the hypothesis of Multicausal
Environmental Severity proposed for sandy beach macrofauna, we believe the classic Intermediate
Disturbance Hypothesis is a more appropriate framework for the meiofauna communities of the studied sites.
© 2011 Elsevier B.V. All rights reserved.
1. Introduction
During the last decades, widespread concern has emerged on
biodiversity loss and its effects on ecosystem processes and
functioning due to the accelerating influence human activities have
on community structure and diversity. As a result, many scientific
articles dealing with biological diversity have been published.
However, in spite of all this effort, most of the published scientific
work has focused on the terrestrial environment (relevant literature
cited in Hooper et al., 2005), while relatively little has been done in
the marine systems. In this context, it is necessary that priorities
should be directed towards inventories of marine species and how
they affect ecosystem services (Snelgrove et al., 1997), the investiga-
tion of biodiversity patterns at large spatial and temporal scales, and
the identification of key mechanisms (Raffaelli, 2006). In 2000, Clarke
and Lidgard (2000) and Gray (2000) pointed out that until that time
there had been almost no studies of beta diversity in the sea and
emphasized the extreme importance such studies would have for the
description of marine biodiversity. Not surprisingly, many subsequent
articles on a variety of marine habitats and organisms have addressed
the issue, or at least incorporated measures of beta diversity (e.g.,
Ellingsen, 2002; Gaertner et al., 2007; Giberto et al., 2007; Koulouri
et al., 2006; Perez-Mendoza et al., 2003; Price, 2002), including a few
focused on meiofauna (De Troch et al., 2001; Veit-Köhler et al., 2010).
While alpha diversity commonly refers to the diversity of a defined
assemblage or habitat (within habitat diversity), beta diversity is
primarily applied as a measure of the change in diversity between
samples along transects, or across environmental gradients, or even at
different spatial configurations or sampling units (Magurran, 2004).
Since beta diversity is not a scale of diversity, Gray (2000) suggested
instead using the term ‘turnover diversity’ as used by Clarke and
Lidgard (2000). However, the term ‘turnover’ usually refers to
temporal changes of diversity and lies at the heart of MacArthur
and Wilson's (1967) theory of island biogeography (Magurran, 2004).
Magurran (2004) stresses that the interplay of alpha and beta
diversity over ecologically and evolutionary time is a topic that
warrants much more consideration. Nevertheless, since the excep-
tional work of Sepkoski (1988) no marine study has examined species
Journal of Sea Research 66 (2011) 205–214
⁎ Corresponding author at: Present address: Hellenic Centre for Marine Research, Institute
of Oceanography, P.O. Box 2214, 71003 Heraklion, Crete, Greece. Tel.: +30 2810 337849.
E-mail address: sevastou@hcmr.gr (K. Sevastou).
1385-1101/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.seares.2011.06.004
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