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Preliminary report of ECOMERS’ 2011 field campaigns
Biodiversity assessment in three alternate stable states of
the biocenosis of photophilic macroalgae of Infralittoral Mediterranean rocky reefs:
Cystoseira forest, Dictyotales- and/or Sphacelariales- dominated assemblages and barren-grounds
Scientists attached to the laboratory Ecosystèmes Marins Côtiers et Réponses aux Stress (ECOMERS)
from the University of Nice Sophia-Antipolis conducted two field campaigns in May and July 2011,
respectively in Corsica and Menorca. The aim was to describe flora and fauna in three different habitats
recognized as alternate stable states of the biocenosis of photophilic macroalgae of Mediterranean rocky
reefs: Cystoseira forest, Dictyotales- and/or Sphacelariales- dominated assemblages and barren-grounds.
Within each habitat, several underwater sampling methods by scuba diving were used for assessing
assemblage structure of (i) sedentary necto-benthic fishes, (ii) cryptic fishes, (iii) vagile macro-invertebrates,
(iv) sessile macro-invertebrates, (v) erect macro-algae and (vi) encrusting macro-algae.
The sampling design included two bioregions (Corsica and Menorca) in order to take into account
natural spatial variability at the scale of the North-Western Mediterranean Basin. In each bioregion, two
localities having all the three different habitats were selected. In Corsica, sampling was done in a protected
locality, “Réserve Naturelle de Scandola”, from 2011 May 7th to 14th
, and in an unprotected locality, “Baie
de Calvi”, from 2011 May 15th
to 21th
. In Menorca, a protected locality, “Reserva Norte de Menorca”, and
an unprotected locality, “Punta Grossa”, were sampled simultaneously from 2011 July 2nd
to 17th
.
This report describes the scientific framework of these field campaigns, the sampling
designs and methods used. Also, it acknowledges the people and organizations who made
these field campaigns possible.
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 2/11
Teams and partners involved
In Corsica, the ECOMERS‟ team was composed of Hazel ARCEO, Adrien
CHEMINEE, Jean-Michel COTTALORDA and Pierre THIRIET. They started at the Marine
Protected Area “Réserve Naturelle de Scandola” which is part of the “Parc Naturel Régional
de Corse”. The park management team headed by Jean Marie DOMINICI kindly hosted the
ECOMERS‟ team. Then, for sampling in “Baie de Calvi”, the ECOMERS‟ team settled at
“Station de Recherches Océanographiques et sous-marines (STARESO)” which is a private
marine station headed by Pierre LEJEUNE. Both the local teams of “Réserve Naturelle de
Scandola” and STARESO provided all logistic support: accommodation, boat and driver,
compressor etc. Moreover, their knowledge was really helpful in choosing the study sites and
appropriately managing weather forecast windows.
In Menorca the ECOMERS‟ team was composed of Adrien CHEMINEE and Pierre
THIRIET and was strengthened by three members of the NGO “Septentrion Environnement”:
Solène BASTHARD-BOGAIN, Médéric BEZELGUES and its director Olivier
BIANCHIMANI. They provided priceless help all throughout the mission, a testament to their
skills, professionalism and involvement into environmental conservation issues. The team was
kindly hosted at the “Estación de Investigación Jaume Ferrer” headed by Joan MORANTA.
This marine station is a branch of the “Instituto Español de Oceanografia”. They provided
boat and accommodation. Moreover, Joan MORANTA and Marta SALES helped in selecting
the study sites through discussions and by accompanying the team during the first field days.
Sampling in the Marine Protected Area “Reserva Norte de Menorca” was allowed by the
“Govern de les illes Balears”.
These field campaigns are part of Pierre THIRIET„s PhD thesis (ECOMERS, 2010-
2013). He is co-supervised by Patrice FRANCOUR and Luisa MANGIALAJO.
Laboratoire Ecosystèmes Marins Côtiers et Réponses aux Stress (ECOMERS)
Pierre THIRIET ([email protected]) & Patrice FRANCOUR (dir. , [email protected])
Université de Nice Sophia-Antipolis, 06108 NICE Cedex 2, France
Association Septentrion Environnement
Olivier BIANCHIMANI (Président, [email protected] )
Maison de la Mer, Anse du Prophète, 244 Corniche Kennedy 13007 Marseille , France
Réserve naturelle de Scandola - Parc naturel régional de Corse
Jean-Marie DOMINICI (Conservateur, [email protected] )
2 rue major Lambroschini BP417 20184 Ajaccio Cedex, France
Station de Recherches Océanographiques et sous-marines (STARESO)
Pierre LEJEUNE (Directeur, [email protected] )
Pointe de la Revellata, BP33 20260 Calvi, France
Estación de Investigación Jaume Ferrer
Joan MORANTA (Coordinador scientifico, [email protected] )
La Mola, Maó, Menorca, Illes Balears, España
Direcció General de Medi Rural i Marí (managing the MPA “Reserva Norte de Menorca”)
Oliver NAVARRO (Coordinador de medio marítimo , [email protected] )
C/ dels Foners, 10 - 07006 Palma, Mallorca, Illes Balears, España
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 3/11
Scientific framework
Human impacts affect benthic communities in direct (e.g. pollution or habitat degradation)
or indirect ways (e.g. overfishing which can potentially induce trophic cascades and related
habitat shift). Coastal rocky shores are among the most threatened ecosystems, as they are
subjected to both land and sea-based impacts. Unfortunately, biodiversity and functioning of
rocky shores ecosystems in the Mediterranean Sea are still very poorly known, and dramatic
changes are happening without us being able to understand or even notice them.
In Infralittoral rocky reefs of the North-Western Mediterranean, areas preserved from
human impact present a biocenosis of photophilic macroalgae characterized by Cystoseira
forests. Cystoseira species are large brown algae that can thrive and reach high densities from
the surface to the limit of the euphotic zone. They are habitat formers and represent an
important habitat for diversified assemblages of associated macroalgae (e.g., sciaphilic
marcoalgae species), invertebrates and fishes. Cystoseira species are very sensitive to direct
and indirect human impacts and their regression has been recorded in several areas of the
Mediterranean Sea.
Indeed, in areas with degraded water quality (e.g. increase in turbidity, waste waters, other
pollutant), Cystoseira forests disappear and are replaced by Dictyotales- , Sphacelariales-
and/or articulated Corallinales- dominated assemblages (forming bush-land). In areas where
date-mussel fishery and/or overfishing (especially of the sea breams) are occuring, they are
replaced by barren grounds, generally characterized by high abundances of herbivores (sea-
urchins). Such barren grounds are almost lacking in primary producers (except encrusting
Corallinales) and can be compared to desertified areas. These new states of marine
ecosystems are generally able to self-maintain and are therefore considered as alternate stable
states.
Macroalgae are primary producers. Moreover, macroalgal assemblages affect the
structural attributes that define a habitat at a fine spatial-scale. These shifts of alternate stable
states (from forests to bush-lands or deserts) may strongly affect biodiversity and ecosystem
functioning. For instance, the structure of macroalgal assemblages affects benthic invertebrate
assemblages, which occupy intermediate trophic levels between primary producers and fish.
Furthemore, more structurally complex habitats may generally provide more shelter for small-
sized fishes reducing their mortality due to predation.
The “trophic” and the “shelter” roles of macroalgae concertedly determine (along with
many others processes such as settlement/recruitment) the patterns of fish distribution and
abundance that are observed in the field. Therefore, in order to disentangle the respective
contribution of these two functions to the whole ecosystem functioning, the necessary
approach appears to be the complementary use of (1) field survey assessing
abundances/biomasses of every trophic level related to each alternate stable states, (2)
stomach contents and isotopic ratios analysis for assessing energy fluxes, and (3) in situ
experiments where conditions are controlled (especially habitat architecture and predation).
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 4/11
Aim of the present study
As the first step in approaching the research problem, the present study is a field survey
aiming to characterize the different alternate stable states in terms of biological and functional
diversity. Specifically, assemblage structures of (i) sedentary necto-benthic fishes, (ii) cryptic
fishes, (iii) vagile macro-invertebrates, (iv) sessile macro-invertebrates, (v) erect macro-algae
and (vi) encrusting macro-algae were assessed.
Definition of the three alternate stable states
(in the framework of the present study)
At the local scale (10m), habitats are characterized by their abiotic features (e.g. depth,
substrate, slope) and biotic features (e.g. macrophytobenthos). In the framework of the present
study which aim to assess macroalgal effect on fish assemblages, abiotic features were fixed.
Each “habitat unit” sampled covered an area of at least 7m in length and 4.5m in width.
The depth was between -4m and -8m. Substrate of the whole area was monolithic rock (as
opposed to blocks, pebbles etc.). Because substrate rugosity is quite difficult to estimate and
therefore to control, steps and crevasses were avoided. Slope was characterized as gentle
(between 0° and 30°). The cover of the habitat unit (7 x 4.5 m²) may appear small but larger
areas fulfilling all these criteria are usually difficult to find because of the small scale
heterogeneity of seascape (fragmentation) in Infralittoral Mediterranean rocky reefs.
The “habitat unit” may belong to any of the three types of habitat defined by biotic
features, i.e. the three alternate stables states:
Forest: at least 80% covered by
Cystoseira spp. (except C.
compressa) forming a canopy that
is quite continuous.
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 5/11
Bush-land: at least 80% covered
by erect macroalgae belonging to
the order Dictyotales (sometimes,
a high component of
Sphacelariaceae family were
considered as acceptable).
Barren-grounds: rock usually
covered by encrusting
macroalgae (mostly Corralinales)
and/or sponges. Cover of erect
macroalgae (usually recruits)
must not exceed 25%.
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 6/11
Sampling design
To describe and compare the three alternate stable states, the sampling design included the
orthogonal fixed factor “Habitat type” encompassing three levels: Cystoseira forest, bush-land
and barren grounds.
To take into account natural spatial variability, the sampling design encompassed 3 nested
spatial scales. At the spatial scale of the North-Western Mediterranean, Corsica and Menorca
(Balearic Islands) were choosen as two bioregions. Within each bioregion two localities were
identified, and within each locality two sites were sampled for all the three types of habitat
(Fig. 1).
Because it could be difficult to find study sites with barren-grounds within No-Take-
Zones (NTZs) and inversely it could be difficult to find study sites with Cystoseira forest
outside NTZs, for each bioregion one locality was sampled within a NTZ (even if it does not
encompass barren-grounds) and the other locality was sampled within a non-protected area
(even if it does not encompas Cystoseira forest). Even if this design might induce a potential
confounding effect between the habitat-type effect and the fishing ban effect, it appeared to be
the only way for achieving comparison of the three habitats. Since this study did not attempt
to assess fishing ban effect (i.e. effectiveness of the NTZ), the non balanced presence of
forest/barren-grounds in NTZ and outside should not be determinant for the interpretation of
results. The factor locality was therefore kept as a random and nested factor within bioregion.
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 7/11
Sampling methods
Five different methods were used for sampling (a) sedentary necto-benthic fishes, (b)
cryptic fishes, (c) vagile macroinvertebrates, (d) erect-macroalgae, and (e) encrusting
macroalgae and sessile macroinvertebrates. Within each habitat unit, one replicate for each
sampling method was performed (See numbers of replicat in Fig. 1).
(a) Sedentary necto-benthic fishes
Highly mobile fish species were not included
because we consider that they are not strongly
associated with any habitat type. They rather
opportunistically exploit resources of habitats they
cross during their daily movement.
Sedentary species assemblages were evaluated
using point count method within a half-disk (9.4m²).
This area was centered within the 7m x 4.5m “habitat
unit” in order to avoid edge effect (Fig. 2). This
method was preferred to transect method because, at
the spatial scale of a standard transect size (25m x
5m), the seascape is usually heterogeneous (multiple
habitats).
One visual census was carried out for each
suitable “habitat unit” (see section above). The diver
kneeled on the floor and waited 1 minute for the fish
to get used to his presence. Then, as fast as possible,
he estimated body size (total length to the nearest cm)
and species of every fish individuals that were present
within the half-disk when estimation started. Data
were recorded using pen and slate. This task usually
lasted less than 3 minutes. The diver then swam
around the sampled area to look for potentially hidden
fish individuals. He also measured the height of the
canopy (if any) in 6 points randomly chosen.
Finally, he put a 1-meter rope on the floor to be
used as a scale when an orthogonal picture of the
whole sampled area was taken. At the laboratory,
macroalgal cover was estimated from the picture (Fig.
3) at the scale of the habitat unit (despite covers are
already ranged by definition of the habitat type, see
section above).
Fig. 2: Position of the diver performing
visual census. The sample area is 9.4m²
and is centered within the habitat unit.
Fig 3. an orthogonal picture
(scaled) of the area sampled.
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 8/11
(b) Cryptic fishes
Visual census methods are not suitable for cryptic fishes as blennioidea. This is especially
true in highly structured habitats such as Cystoseira forests because small fishes may be
hidden within the canopy. One suitable method for sampling cryptic fishes is the use of an
“enclosed quinaldine station”.
In order to delineate the sample area and to prevent fish from escaping, a mesh-fence was
used. It was a cylinder 112.8cm in diameter (1m²) and 100cm in height, top- and bottom-less.
It was laid vertically on the seafloor (Fig. 4a). Quinaldine was sprayed through the canopy to
put fish to sleep (Fig. 4b). Fish were then collected using an airlift pump (Figure 4c).
All fish gathered
in the net bag of the
airlift pump were
fixed in 70%
alcoholic solution.
At the laboratory,
the specimens were
measured (total
length to the nearest
mm) and weighed
(to the nearest g)
(Fig. 5).
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 9/11
(c) Vagile macroinvertebrates
The mesh size of the airlift pump
collector (the bag filtering the output, Fig. 6)
is 1mm. Therefore, vagile macroinvertebrates
were collected at the same when pumping
cryptic fishes. Some macroalgae and some
detritus are also pumped. The same day of the
collection, invertebrates specimens were
extracted from the bag content and were
stored in 70% alcoholic solution. Sample
sorting and quantification will be done in the
next months.
Fig. 6: A diver closing the collector (1mm
mesh-size) as soon as the pump was turned off.
(d) Erect macroalgae
A 625cm² quadrat was randomly positioned.
Every erect macroalgae was scraped using chisel
and hammer. It is worth noting that some
epiphytic-macroinvertebrates (e.g. sponges on
Cystoseira. spp) could be part of the sample.
Sample of erect macroalgae were processed
within the same day of their collection.
Individuals were sorted and weighed (fresh)
according to operational taxonomic units (Fig. 7).
The genus Cystoseira and the order Dictyotales
(eventually also the family Stypocaulaceae) were
identified up to the species level. In case of doubt
about the species, some samples were dried in herbarium for later identification. All other
individuals were pooled into wide operational taxonomic units and weighed. Since the aim of
the study is to characterize biotic features of fish habitat, simple functional groups were
therefore preferred.
(e) Encrusting macroalgae and sessile macroinvertebrates
After collecting erect macroalgae, an orthogonal picture of each quadrat was taken. Cover
of encrusting coralline algae, encrusting non-coralline algae, sponges, other sessile
invertebrates and bare substrate will be estimated later based on the orthogonal picture(Fig. 8)
Fig. 7: Macroalgae pooled in operational
taxonomic units
Fig. 8 : orthogonal
pictures of quadrats after
scraping in Cystoseira
forest, Bush-land and
Barren-grounds (from
left to right).
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 10/11
Summary of data collected and future analysis
For each trophic level, assemblage structure will be compared between habitats using the
different metrics available both in multivariate and univariate contexts. Within each habitat,
correlation between assemblage structure of different trophic level will be explored (e.g. in
Cystoseira forest, correlation between fish density and weight/height/cover of Cystoseira
brachycarpa var. balearica).
Table summarizing all the metrics measured during the study. A total of 24 sites were
sampled. Number of replicat is the number of measures per site.
Trophic level Metrics available
Number
of
replicats
Sedentary
demersal fishes
Body-size and density (indiv./9.4m²) per species
(biomass may be calculated using existing weight-length
relationships)
6
Cryptic fishes Body-size, body-weight and density (indiv./1m²) 3
Vagile
macroinvertebrates
Total weight of specimens (from 625cm²) pooled according
to operational taxonomic units 3
Sessile
macroinvertebrates
Covers (out of 625 cm²) of each operational taxonomic
units (biomass may be calculated using existing weight-
area relationships)
3
Erect macroalgae
Total weight of specimens (from 625cm²) pooled according
to operational taxonomic units 3
Height of the canopy 36
Covers (out of 7 x 4.5 m²) of Cystoseira forest, Bush-land
and barren-grounds 6
Encrusting
macroalgae
Covers (out of 625 cm²) of each operational taxonomic
units (biomass may be calculated using existing weight-
area relationships)
3
Scientific knowledge and management advices expected
The expected results of this study will allow quantifying the loss of
biological and functional diversity due to the regression of Cystoseira forests. At
date, only few studies pointed out Cystoseira spp. as key species. The outcome
of the study might be used as additional argument for establishing more drastic
protection/management measures of the coastal marine environment. Moreover,
analysis of distribution patterns is the first step toward a better understanding of
the mechanisms involved in the self-maintaining of the altered stable states
(Bush-land or Barren-grounds). In perspective, ecological engineering methods
might be designed to facilitate the recovery of Cystoseira forest.
Preliminary report of ECOMERS’s 2011 field campaigns – [email protected] 11/11
Concluding remarks
Most of data and biological samples collected during these field campaigns
are still to be analyzed. Statistical analyses and ecological interpretation are not
completed. This will be the subject of a second report.
We wish to thank all the scientists, divers and partners cited above who made
these campaigns possible, and especially for making everything go smoothly.
Divers from the ECOMERS‟ team with (a) Jean-Marie DOMINICI on the boat
of Scandolla Reserve, (b) Alexandre VOLPON on the boat of STARESO, and (c)
Marta SALES on the boat of the Estación de Investigación Jaume Ferrer.