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Benthic survey of natural and artificial reefs off Mar del Plata, Argentina, southwestern Atlantic/Estudios del bentos de arrecifes naturales y artificiales de Mar del Plata, Argentina, Atlantico sudoccidental.

INTRODUCTION

During the past years, the fishing effort has increased and coastal resources had been more heavily impacted due to trawling in Buenos Aires (Bertolotti et al., 2001, 2008). Bottom-trawl fisheries disturb benthic communities and habitats and can reduce biomass, production and diversity, which at the end may change ecosystem properties (Collie et al., 2000; Kaiser et al., 2000; Hiddink et al., 2006a, 2006b). Within this context, natural reefs (NR), a strip or ridge of rocks or coral that rises to or near the surface of the water, provide refuge to coastal fishes and habitat to numerous invertebrates and algae as they cannot be disturbed by demersal fishing activities. Accordingly, artificial reefs (AR) have been suggested as a tool for the conservation of biodiversity, mitigation and rehabilitation of habitats under anthropogenic impacts (Perkol-Finkel & Benayahu, 2005, 2007; Perkol-Finkel et al., 2006).

An artificial reef is one or more objects of natural or human origin deployed purposefully on the seafloor to influence physical, biological, or socio-economic processes related to living marine resources (Seaman, 2000). Accidental shipwrecks have been also classified at times as AR. This definitions reflects hypotheses that the ecological processes of artificial reefs are (or should be) functionally equivalent to those of natural benthic systems in an area (Seaman, 2000). If the last is true, after a long term period of submersion we could expect that ARs and NRs will have a similar community structure (Perkol-Finkel et al., 2006). On the contrary, if the AR is not mimicking the NR in a proper way, the resulting benthic communities should differ (Glasby & Connel, 2001).

The use of AR to mitigate the effects of human activities has been stressed out by fisheries managers and conservation biologists. However, there is still a debate regarding their performance in attracting fish resources from the surroundings and depleting them or enhancing production in the whole area, otherwise lost (Carr & Hixon, 1997; Grossman et al., 1997; Svane & Petersen, 2001). In the case of epibenthic organisms, the introduction of an artificial habitat clearly increases the local production of fouling assemblages; its structural complexity and submersion period affect the colonizing community (Baine, 2001; Svane & Petersen 2001; Perkol-Finkel et al., 2006).

Mar del Plata coastal area is characterized by numerous rocky outcrops that extend from the intertidal to several miles seawards, up to 25 m depth. The outcrops and surrounding areas are subjected to many human activities, including commercial and recreational fisheries and diving. In addition, some shipwrecks have built extended AR during the past decades, which provide hard substrata suitable for the recruitment of benthic and refuge for fish species. The fauna and habitats remain largely unknown in the study area; only some cnidarian groups were surveyed (Genzano et al., 2002; Excoffon et al., 2004). On the other hand, diving at Mar del Plata outcrops is a more recent focus for the diving centers of the region, who have proposed the creation of submarine parks to boost diving activities in these almost unknown marine communities.

In this paper we study the benthic communities of coastal outcrops off Mar del Plata (both natural and artificial reefs). The main goals are 1) to describe for the first time the macrofaunal assemblages of subtidal hard bottoms, 2) to compare the macrobenthic assemblages settled on shipwrecks with nearby outcrops to determine the effectiveness in reflecting the natural communities of the region, after a long-term period of submersion (60 years), 3) to characterize the fish fauna associated with NRs and ARs, and 4) to obtain reliable information to management and conservation of these coastal habitats.

Buenos Aires coast comprises sandy beaches and numerous isolated intertidal hard bottoms of loess (consolidated sedimentary rocks, Pleistocene). The coast off Mar del Plata city is exceptionally different due to the presence of quartzite rocks originated in the Tandilia mountainous system (SE of Buenos Aires Province), during the Ordovician, which extends several miles seawards up to 25 m depth (Genzano, 2010). The sublittoral rocky area is estimated at ca. 14 [km.sup.2] and constitutes the largest hard substrate in the zone. Although the area is indicated as continuous in the nautical charts, it is actually a group of isolate outcrops of variable size surrounded by sand and shell debris and with different physiognomy and faunal composition. The outcrops are well-known to divers and fishermen, which commonly named them Restingas (outcrops up to 12-14 m depth) and Bancos (outcrops 18-23 m depth) (Fig. 1). These outcrops are conformed by big fragmented quartzite boulders, with the exception of a sedimentary slope originated during the Quaternary period (Neozoic era) which is located over the ortoquarcitic rocky floor in Banco de Afuera. This slope is 1-1.5 m height conformed by a more fragile consolidated sedimentary rock (coquina with abundant shell fragments).

The area is affected by a littoral current (south to north), semi-diurnal tidal currents and high-energy waves; storms and winds from SE are frequent, mainly in autumn and winter (Lanfredi et al., 1992). During this period big amounts of sand and fragments of mollusk valves accumulate in depressed areas and crevices. Storms and winds notably produce sediment re-suspension. Consequently, diving is usually developed under dangerous conditions, with water visibility ranging between 0.5-6 m, average 3 m. Water temperature is variable through the year, maximum summer values reach 18-19[degrees]C and minimum winter values of 7-8[degrees]C were recorded.

Six vessels sank in the area more than 50 years ago, which also denotes the difficult conditions for nautical activities. The shipwrecks of the freighter James Clunies (149 m length, sunk in 1949 at 10 m depth), over a restinga are frequently visited by local divers. In addition, in 1981 a submarine park was built on soft bottoms close to Restingas and Bancos, by sinking the wood-vessel Cristo Rey (45 m length) at 22 m depth, thus creating another site of interest for divers (Inurieta, 2005; Genzano, 2010).

MATERIALS AND METHODS

Several exploratory dive surveys were conducted between July 2000 and May 2005 in the study area with the aim to provide an assessment of the habitat types and to select sampling sites. All census and collections were carried out by SCUBA diving at the different rocky outcrops and artificial reefs (shipwrecks), between 1 and 23 m depth. A total of 26 dives ([approximately equal to]900 min of immersion) were performed during the study. Topographical characterization and macro-invertebrates census were performed during November and December 2005, austral spring (a period in which most species would likely be present) in 11 sites: Restingas Faro (R0), Primera (R1), Segunda (R2) and Tercera (R3), Bancos Pescadores Sur (PS), del Medio (BM), Banco de Afuera (BA) and the sedimentary slope in BA (BAS) and shipwrecks James Clunies vertical (JCV) and horizontal (JCH) substrata and Cristo Rey (CR) (Fig. 1).

In each site three parallel 10 m chains, horizontally positioned, separated by 5 m were laid out at random, performing line-intercept transects (Loya, 1978), writing down the nature of the substrate (coverage over the length on the chain), according with predefined categories of substrata (modified from Woodman, 1999): boulders (human size or bigger), rocks (size of an arm or legs), stones (hand size), pebbles (finger size), mud/sand/bioclast (fine particle). Live cover of conspicuous macrobenthic organisms (algae, sponges, cnidarians, blue mussels, sea stars, sea urchins) was also measured. These data provided information on the relative cover of major benthic groups, used to define community types. Coverage was recorded counting the number of chain links covering the different species, method previously used in cnidarian's censuses in the area (Genzano et al., 2002).

In addition, qualitative samples were collected in some particular microhabitats to identify taxa when stereoscopic microscope observation was needed in order to give the baseline inventory list from this almost unexplored area, which was complemented by underwater observations and video recording conducted during 2007 and 2008 using rovers diver transects (RDT).

As a measure of topographic complexity, the Rugosity index (Ri) was calculated using the ratio of a length of the chain moulded to the surface to the linear distance between its start and end points and it was expressed as percentages as follows: Ri (%) = (1-[length of chain in situ / total length of chain]) x 100 (Luckhurst & Luckhurst, 1978; Francour & Koukouras, 2000). In addition, the Cavity index (Cav), was calculated as follows: Cav = [B.sup.2] * 100/([A.sup.2]+[B.sup.2]), where "A" is the mean size of the external face of blocks and "B" is the mean size of cavity apertures between blocks and rocks (Ruitton et al., 2000). Average height, the distance from the bottom substrate to the top of the 10 highest blocks within a transect, was also estimated as other assessment of habitat architecture (Gratwicke & Speight, 2005).

Quantification of fishes represent a special problem because of their mobility, hence censuses are necessary subjective and reliant upon water condition. Visual censuses were performed swimming freely throughout a dive site during 15 min in each survey and recording every fish species that could be positively identified (Lessios, 1996). Since local species do not conform dense fish schools, each recorded species was assigned into one of four relative abundance categories (modified from Pattengill-Semmens, 2001): occasional (O): a single individual, scarce (S): 2-5 individuals, abundant (A): 6 a 10 and very abundant (Va): more than 10 individuals observed during the immersion. Frequency was calculated as the ratio between numbers of dives in which a species was found and the total dives conducted. Three categories were assigned: less frequent (LF): recorded in less than 20%; frequent (F): recorded between 20-50% and very frequent (VF): recorded in more of the 50% of total dives.

[FIGURE 1 OMITTED]

A non-parametric multivariate analysis was applied to environmental and biological data. Similarity of sites using physical variables (depth, substrate categories and topographic complexity) was determined by ordination methods (MDS, using group average sorting of the Euclidean distance of physical data). In a similar way, changes in species composition among sampling sites was determined by ordination methods (MDS, group average sorting of the Bray-Curtis similarity measures on cover data) using PRIMER software (Clarke & Warwick, 2001). The SIMPER analysis (similarity percentages) was used to identify those species which contributed most to (dis)similarities among and within groups. This analysis calculates the "average similarity" (contribution of the [i.sub.th] species to the overall dissimilarity between the groups considered) and the "internal similarity" (contribution each species makes to the average similarity within each group considered) (see Clarke & Warwick, 2001 for details). The ANOSIM test was used to search for differences between groups of sampling sites (R0 to CR: natural vs artificial reefs, horizontal vs. vertical communities). This permutation test analyses composition differences between replicates within sites contrasted with differences between sites, computing an R-statistic under the null hypothesis "no differences between sites". R falls between -1 and 1, so R is ~0 if the null hypothesis is true and R = 1 if all replicates within sites are more similar to each other than are replicates from different sites. Finally, the BIO-ENV procedure was used to determine which set of environmental variables (similarity calculated with the Euclidean distance coefficient) best explains the biological matrices (presence-absence data, using Bray-Curtis similarity measure) (Clarke & Warwick, 2001).

RESULTS

The first sublittoral Rocky Outcrop (R0) is located 300 m off the costal line; its shallower zone remains partially exposed to the air during extraordinary low tide. This outcrop is conformed mainly by boulders and big rocks which reach 1-2 m depth. Other two outcrops of similar topography (R1 and R2) are found between 5-10 m depth. Conform the depth increases, rocks and stone coverage are more important and, consequently seafloor roughness is higher. Broken mollusk shells and sand are usually accumulated in cavities, crevices and channels. The outcrop R3 lies between 12-14 m depth and BM, BA and PS between 19-23 m depth. The most remarkable difference in topography and composition of the sea floor is found in BA, due to the presence of a slope 1-1.5 m height (BAS) of consolidated sedimentary rocks (coquina with abundant shell fragments).

Two shipwrecks constitute extensive ARs. The first one, the vessel James Clunies, is located on rocky bottoms in R2; the boiler remains in a vertical position up to 2 m from the seafloor (JCV), while the deck offers horizontal substrata (JCH). The other AR, the vessel Cristo Rey (CR), is located in soft bottoms near R3, offers numerous and discontinuous hard substrata of complex topography (Table 1).

A total of 108 taxa were recorded in the study area, including 7 macroalgae, 98 invertebrates and 3 tunicates. Twenty one were the most conspicuous taxa both NR and AR and they were used in quantitative assessment of living cover of macrobenthic organisms (see above). The remainder species correspond mainly to small size organisms associated to mussel beds and/or epizoites on big clumps hydroid colonies. Other macrobenthic organisms were rarely observed due to their difficult detection because of scarce abundance or cryptic habits (Tables 2 and 3).

The pattern of samples agrees with depth and position in the substrate (Fig. 2). A gradient from substrates exposed during exceptional low tides to deepest sites was observed. The outcrop R0, dominated by Corallina officinalis and Ulva lactuca (72 and 24% cumulative contribution respectively, SIMPER analysis), was clearly separated from the rest of the sampling sites, which followed a gradient from shallow to deep ones: R1, R2, R3, JCH, PS, BA (typical species Mytilus platensis, 69.0-99.2%), CR, BM and BAS (Tripalea clavaria 57.6-98.3%) (Fig. 3). No differences were found between horizontal AR and the adjacent NRs. Main differences were found between vertical vs. horizontal communities, both in NR and AR (BAS and JVC respectively) (Fig. 3, see Tables 4 and 5). JCV was slightly separated from the rest due to its distinct biological composition established in a vertical artificial substrate (Anthothoe chilensis 99.3%) (Fig. 3).

The best combination of structural features explaining the biological matrix (BIO-ENV) included the combination of the following variables: Fine Sediment, RI, Height and Depth (Sample statistic Rho: 0.495); RI, Height, Depth (Sample statistic Rho: 0.493) and Height and Depth (Sample statistic Rho: 0.490).

Fifteen reef fish species were recorded during the 35 visual censuses conducted during this study. Species richness was lower in shallower outcrops than in deeper ones (8 vs. 14 species). Six species were found in all AR and NR (Table 6). Overall, only 4 species (26.7% of the total) were classified as frequent or very frequent: Diplodocus argenteus, Pagrus pagrus, Dules auriga and Pinguipes brasilianus, the last one was the most abundant in all sites. Acanthistius brasilianus was recorded with highest abundance and frequency values in the shipwrecks (AR) but infrequent in the outcrops (NR), with the exception of caves and crevices from BAS. The few specimens of Pseudopercis semifasciata observed during the visual censuses were reported in the same cryptic habitats (AR and BAS). The remaining fish species occurred with low frequency, usually single specimens, with the exception of a large shoal of Seriola lalandei observed in the deep outcrops.

DISCUSSION

Our results provide information on sublittoral faunal assemblages from unexplored locations of southwestern Atlantic coasts. The rocky outcrops in the area of Mar del Plata were composed by distinct benthic communities than those previously studied from nearby rocky intertidal and soft bottoms areas. The intertidal zones are dominated by the mytilid Brachidontes rodriguezi (Olivier et al., 1966; Penchaszadeh, 1973; Scelzo et al., 1996). On the other hand, subtidal coarse sandy adjacent areas between 18 and 25 m depth are dominated by the sea urchin Encope emarginata, while deeper and finer bottoms up to 64 m depth show the presence of Mytilus platensis banks of high species richness (Bremec & Roux, 1997). In the rocky outcrops herein studied, red calcareous algae dominated the shallowest NR while a conspicuous mytilid assemblage was present in NRs deeper than 3 m; sea anemones were more abundant between 6-10 m and soft-corals at nearly 20 m depth (Fig. 4). Epizoism was an important phenomenon in the area, mainly on sertulariid and plumulariid hydroids (see Table 1).

[FIGURE 2 OMITTED]

A wide array of structural features is known to affect the settlement of benthic invertebrates onto natural and artificial substratum, including spatial orientation, structural complexity, substratum composition and texture (Moura et al., 2006; Perkol-Finkel et al., 2006; Thanner et al., 2006). Community structures developed in horizontal AR (JCH) after nearly sixty years were similar to neighboring NR communities (R1 and R2), while dominant taxa differed notably within vertical or horizontal ARs and NRs. Perkol-Finkel et al. (2006) found that if submersion time is prolonged, ARs can mimic adjacent NRs only if they possess similar structural features. However, if they differ structurally, their communities will remain distinct. Our results are in accordance with previous works that demonstrated clear consistent differences, even considering long periods of AR submersion, between epibiotic assemblages settled on horizontal and vertical surfaces (Moura et al., 2006; Perkol-Finkel et al., 2006; Thanner et al., 2006). Proximity to the seafloor and shading can markedly influence the development of distinct epibiotic assemblages in subtropical and tropical waters (Glasby & Connell, 2001; Perkol-Finkel et al, 2006).

[FIGURE 3 OMITTED]

Suspended particles are highly dependent on the existence of elevated structures mostly because the water current velocity increases when passing by the edge of them (Falcao et al., 2009). The a zoo-xanthellate soft-coral Tripalea clavaria, which also occur in soft bottom up to 70 m depth, depends on the flow current to feed (Olivier et. al., 1968; Acuna et al., 2004). Thus, its abundance in BAS and CR could be explained by the vertical disposition of these substrata that affect the fluxes of suspended material. The small sea anemone Anthothoe chilensis monopolized the available artificial vertical substrata, i.e. 5000 ind [m.sup.-2] in JCV (Fig. 5). Such high abundance could be explained by its preference to colonize substrata where water currents guarantee a continuous supply of food and avoid the negative effects of sedimentation. This species also occurs in dense patches in the intertidal fringe (Excoffon et al., 1997). Its higher abundances could be also favored by its reproductive strategy by longitudinal fission (Excoffon & Acuna, 1998). Other fast growing sea anemone, Metridium senile (Linnaeus, 1761), has a similar asexual mechanism and is usually found colonizing shipwrecks in other coastal temperate waters in Belgium (Zintzen et al., 2008) and Patagonia (G. Genzano, pers. obs.).

In AR, spatial complexity increases habitat heterogeneity, providing increased areas of refuge for fish populations from predation and competition. Reef fishes also represent an attraction for recreational divers, and for sport angler fishers (Galvan et al., 2009). Fifteen species were recorded in the analyzed area. Abundance and frequency values registered by visual censuses for some species could be underestimated due to their particular behavior; e.g. Dules auriga and Bovichthys argentinus are two shy fishes which usually avoid divers and Triathalassothia argentina is a cryptic and sedentary species with nocturnal habits. However; the data clearly indicated that species having strong site fidelity, i.e. Acanthistius brasilianus (Irigoyen & Venerus, 2008) and Pseudopercis semifasciata, were clearly more abundant and/or frequent in ARs (shipwrecks) compared to the surrounding NRs, suggesting a preference for habitat with more complex structure. In NRs these species occur only in cliffs and crevices of BAS but they are rare or absent in the less structurally complex outcrops.

This pioneer study in the surveyed area clearly shows that outcrops allow the settlement of diverse and rich assemblages. Artificial reefs allow not only the settlement of benthic invertebrates but also provide refuge for fishes. As recreational fishery and diving are intensifying its activities in Mar del Plata, where tourism is one of the main economical resources, the protection of such areas should be considered in conservation plans. Future management plans should have into account the frequency of dive-expeditions in these coastal habitats.

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DOI: 10.3856/vol39-issue3-fulltext-15

ACKNOWLEDGEMENTS

We acknowledge CASE Diving Center for their help during field work and logistical support. This paper was funded by PICT 2007 02200 and 2008 01194; EXA 460/09, CEN 031/10. This is INIDEP contribution 1690

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Received: 22 December 2010; Accepted: 11 October 2011

Gabriel Genzano (1,2), Diego Giberto (2,3) & Claudia Bremec (2,3)

(1) Estacion Costera Nagera, Departamento de Ciencias Marinas, Facultad de Ciencias Exactas y Naturales Universidad Nacional de Mar del Plata, Funes 3350, Mar del Plata, Argentina

(2) Instituto de Investigaciones Marinas y Costeras (IIM y C), Universidad Nacional de Mar del Plata Consejo Nacional de Investigaciones Cientificas y Tecnicas, Argentina

(3) Proyecto Ecologia Pesquera, Laboratorio de Bentos, Instituto Nacional de Investigacion y Desarrollo Pesquero (INIDEP), Paseo V.O. No. 1, Mar del Plata, Argentina

Corresponding author: Gabriel Genzano (genzano@mdp.edu.ar)
Table 1. Measures of substrate topography (average [+ or -]
standard deviation) along the natural and artificial reefs of the
study area. Natural reefs: Restingas Faro (RO), Primera (R1),
Segunda (R2) and Tercera (R3), Banco Pescadores Sur (PS), Banco del
Medio (BM), Banco de Afuera (BA) and the sedimentary slope in BA
(BAS). Artificial reefs: James Clunies horizontal (JCH) and
vertical (JCV), and Cristo Rey (CR). RI: rugosity index. CI: Cavity
index.

Tabla 1. Medidas de la topografia del sustrato (promedio [+ or -]
desviacion estandar) en los arrecifes naturales y artificiales del
area de estudio. Arrecifes naturales: Restingas Faro (RO), Primera
(R1), Segunda (R2) y Tercera (R3), Banco Pescadores Sur (PS), Banco
del Medio (BM), Banco de Afuera (BA) y cordon sedimentario en BA
(BAS). Arrecifes artificiales: James Clunies horizontal (JCH) y
vertical (JCV), y Cristo Rey (CR). RI: indice de rugosidad. CI:
indice de cavidad.

                            Natural reef

                        RO                  R1

Boulder (%)      46.O [+ or -] 7.5   67.5 [+ or -] 5.5
Rocks (%)        37.O [+ or -] 5.5   26.5 [+ or -] 4.7
Stones (%)        8.7 [+ or -] 2.5    7.2 [+ or -] 1.7
Pebbles (%)              O            O.3 [+ or -] O.2
Fine Sediments    8.7 [+ or -] 1.6    2.7 [+ or -] O.8
  (%)
RI (%)           23.3 [+ or -] 2.5   42.8 [+ or -] 2.1
CI (%)                 22.25               19.16
Height (cm)      16,1 [+ or -] O.7     2O [+ or -] 1.O
Depth (m)                1                   7

                            Natural reef

                        R2                  R3

Boulder (%)      42.3 [+ or -] 7.4    3.O [+ or -] 1.4
Rocks (%)        27.3 [+ or -] 4.8   55.O [+ or -] 2.9
Stones (%)       16.7 [+ or -] 2.8   13.6 [+ or -] 2.3
Pebbles (%)       1.3 [+ or -] O.6    2.8 [+ or -] O.9
Fine Sediments   11.3 [+ or -] 2.1   24,9 [+ or -] 2,6
  (%)
RI (%)           35.8 [+ or -] 1.2   44.5 [+ or -] O.3
CI (%)                 27.69               36.4
Height (cm)      18,1 [+ or -] 1.2   27,7 [+ or -] 3.O
Depth (m)               1O                  14

                             Natural reef

                        BM                   BA

Boulder (%)       8.7 [+ or -] 3.3   1.67 [+ or -] 1.7
Rocks (%)        45.O [+ or -] 5.5   46.9 [+ or -] 4.9
Stones (%)       21.3 [+ or -] 3.6   21.3 [+ or -] 3.5
Pebbles (%)       2.7 [+ or -] O.9   1.33 [+ or -] O.63
Fine Sediments   19,7 [+ or -] 2,9   29,7 [+ or -] 2,8
  (%)
RI (%)           49.6 [+ or -] 1.4   39.4 [+ or -] 1.3
CI (%)                 41.56                41.4
Height (cm)      35,O [+ or -] 3.5   24,O [+ or -] 2.3
Depth (m)               2O                   21

                             Natural reef

                        BAS                  PS

Boulder (%)      85.O [+ or -] 2.5    12.3 [+ or -] 4.5
Rocks (%)         5.3 [+ or -] 1.8    38.7 [+ or -] 4.9
Stones (%)        1.O [+ or -] O.6    17.3 [+ or -] 3.7
Pebbles (%)       O.3 [+ or -] O.3    1O.3 [+ or -] 2.9
Fine Sediments    8,3 [+ or -] 1.8    36.O [+ or -] 3.1
  (%)
RI (%)              59 [+ or -] O.65  35.7 [+ or -] 1.6
CI (%)                   O                  44.3
Height (cm)       6O,O [+ or -] 1O.O  2O,O [+ or -] 3.8
Depth (m)                22                  21

                               Artificial reef

                        JCH          JCV          CR

Boulder (%)              O            O            O
Rocks (%)                O            O    16.5 [+ or -] 5.9
Stones (%)               O            O    13.3 [+ or -] 5.3
Pebbles (%)              O            O     1.4 [+ or -] O.8
Fine Sediments   5.5 [+ or -] 3.O     O    34.1 [+ or -] 7.3
  (%)
RI (%)                   O            O    51.9 [+ or -] 3.3
CI (%)                   O            O            O
Height (cm)      1O,O [+ or -] 2.2   2OO   6O,O [+ or -] 8.3
Depth (m)               1O           1O           23

Table 2. List of the taxa observed at sublittoral natural
outcrops andartificial reefs of the study area. (*) Taxa used in
quantitative analysis, 1) Epizoite on bigger hydroid colonies,
2) ecto-parasites on the hydroid Amphisbetia operculata,
3) endolithic organisms inhabiting BAS; 4) endobiont in the the
gorgonean Tripalea clavaria, 5) specimens inhabiting
fine sediments accumulated in cavities and crevices.

Tabla 2. Lista de taxa observados en los arrecifes naturales y
artificiales del area de estudio. (*) Taxa utilizados en los
analisis cuantitativos, 1) Epizoicos sobre grandes colonias
hidroides, 2) ecto-parasitos sobre el hidroide Amphisbetia
operculata, 3) organismos endoliticos que habitan el BAS,
4) endobiontes en el gorgonaceo Tripalea clavaria; 5) especimenes
que habitan los sedimentos finos acumulados en cavidades y grietas.

Algae:           Ceramium virgatum Roth 1797; Codium fragile
                 (Suringar); Corallina officinalis L(*)
                 Polysiphonia sp.(*) Pterosophonia sp.;
                 Rhodymenia sp.(*); Ulva lactuca L.(*).

Porifera:        Callyspongia sp.; species indet. species
                 indet. 2(*), species indet. 3(*), species
                 indet. 4(*), species indet. 5(*), species
                 indet. 6(*).

Cnidaria:        Aglaophenia acacia Allman, 1883;
                 Amphisbetia operculata (Linnaeus, 1758)(*);
                 Antholoba achates (Drayton, 1846);
                 Anthothoe chilensis (Lesson, 183O)(*);
                 Astrangia rathbuni Vaughan, 19O6;
                 Aulactinia sp.(*); Bimera vestita Wright,
                 1859; Campanularia agas Cornelius, 1982(1);
                 Choriactis laevis (Carlgren, 1899); Clytia
                 gracilis (Sars, 1851)(1); Corynactis carnea
                 Studert, 1878(*); Coryne eximia Allman,
                 1859; Dynamena cornicina McCrady, 1859;
                 Ectopleura crocea (Linnaeus, 1758);
                 Familia Plexauridae species indit.;
                 Filellum sp.(1) ; Halecium beanii
                 (Johnston, 1838); Halecium delicatulum
                 Coughtrey, 1876; Hebella sbandens
                 (Bale, 1888)(1); Monostaechas quadridens
                 (McCrady, 1859); Monotheca pulchella
                 (Bale, 1882); Obelia sp., Order
                 Ceriantharia species indet.; Plumularia
                 setacea (Linnaeus, 1758)(*); Sagartia
                 troglodytes Price in Johnston, 1847;
                 Sertularella mediterranea Hartlaub,
                 19O1(*), Sertularella striata Stechow,
                 1923; Tricnidactis errans (Pires, 1988);
                 Tripalea clavaria (Studert, 1878)

Plathelminta:    Notoplana sp.

Nematoda:        Deontostoma sp., Enoplus sp.,
                 Pseudocella sp.

Nemertina:       species indet. 1

Bryozoa:         Aetea anguina (Linnaeus, 1758)(1);
                 Bicelariella sp.; Bugula sp.; Celleporella
                 sp., Membranipora sp.; Osthimosia sp.(1);
                 species indet.

Crustacea:       Balanus improvisus (Darwin, 1854); Balanus
                 trigonus (Darwin, 1854); Caprella equilibra
                 Say, 1818; Corophium insidiosum (Crawford,
                 1937); Jassa falcata (Montagu, 18O8);
                 Halicarcinus planatus (Fabricius, 1775);
                 Idotea baltica (Pallas, 1772); Leucipa
                 pentogona Milne Edwards, 1833; Orden
                 Tanaidacea species indet.; Pachycheles
                 laevidactylus Ortmann, 1892; Platyxanthus
                 crenulatus (Milne Edwards, 1879);
                 Pleoticus muelleri (Bate, 1888);
                 Serolis sp.

Annelida:        Capitela "capitata" sp.; Diopatra viridis
                 Kinberg 1865; Eunice sp.; Halosydnella
                 australis (Kinberg, 1855); Lumbrineridae
                 species indet.; Neanthes succinea (Frey &
                 Leuckart, 1847); Nereidae species indet.;
                 Ophelidae species indet.; Orbiniidae
                 species indet.; Phyllochaetopterus socialis
                 Claparede, 187O; Polonoidae species indet.;
                 Polygordius appendiculatus Frainpont, 1887;
                 Procerastea halleziana Malaquin, 1893(2);
                 Serpulidae species indet.; Syllis sp.;
                 Terebridae species indet.

Sipunculida:     Themistes petricola (Amor, 1964)(3).

Mollusca:        Aequipecten tehuelchus (d'Orbigny, 1846);
                 Anachis sp.; Atrina seminuda (Lamarck,
                 1819); Calliostoma sp.; Crepidula sp.;
                 Chaetopleura sp.; Doridoidea species
                 indet.; Fisurellidea megatrema d'Orbigny,
                 1841; Litophaga patagonica (D'Orbigny,
                 1847)(3); Musculus viator (d'Orbigny,
                 1846)(4); Mytilus platensis D'Orbigny,
                 1846(*); Ostreapuelchana D'Orbigny, 1841;
                 Tegula sp.

Pycnogonida:     Achelia assimilis (Haswell, 1885);
                 Tanystilum orbiculare Wilson, 1878.

Echinodermata:   Arbacia dufresnei (Blainville, 1825)(*);
                 Asterina sterifer (Mobius, 1859)(*);
                 Dendrochirotida species indet.(4),
                 Ophioplocus januarii (Lutken, 1856).

Tunicata:        species indet.; Molgula sp.; Sycozoa
                 umbellata (Michaelsen, 1898).

Table 3. Living cover (average [+ or -] standard deviation)
of macrobenthic organisms at natural and artificial reefs
of the study area.

Tabla 3. Cobertura (promedio [+ or -] desviacion estandar)
de organismos macrobentonicos en arrecifes naturales y
artificiales del area de estudio.

                                Natural reef

                            R0                   R1

Ulva lactuca          9.0 [+ or -] 2.5           -
Corallina            26.0 [+ or -] 5.8           -
  officinalis
Polysiphonia sp.             -           0.5 [+ or -] 0.2
Rhodymenia sp.               -           0.2 [+ or -] 0.1
Porifera sp. 1               -           2.6 [+ or -] 1.4
Porifera sp. 2               -                   -
Porifera sp. 3               -                   -
Porifera sp. 4               -                   -
Porifera sp.5                -                   -
Porifera sp. 6               -                   -
Amphisbetia                  -                   -
  operculata
Anthothoe                    -           0.17 [+ or -] 0.1
  chilensis
Aulactinia sp.               -            0.1 [+ or -] 0.05
Corynactis carnea            -                   -
Ectopleura crocea            -                   -
Sertularella                 -                   -
  mediterranea
Plumularia setacea           -                   -
Tripalea clavaria            -                   -
Mytilus platensis    9.7 [+ or -] 4.0    12.7 [+ or -] 4.2
Arbacia dufresnei            -            0.1 [+ or -] 0.05
Asterina sterifer            -            0.1 [+ or -] 0.06

                                 Natural reef

                             R2                   R3

Ulva lactuca                 -                     -
Corallina                    -                     -
  officinalis
Polysiphonia sp.      0.1 [+ or -] 0.09            -
Rhodymenia sp.               -             1.4 [+ or -] 0.2
Porifera sp. 1               -                     -
Porifera sp. 2               -                     -
Porifera sp. 3               -                     -
Porifera sp. 4               -                     -
Porifera sp.5                -                     -
Porifera sp. 6               -                     -
Amphisbetia                  -             0.1 [+ or -] 0.1
  operculata
Anthothoe             1.8 [+ or -] 0.5     1.9 [+ or -] 0.6
  chilensis
Aulactinia sp.               -                     -
Corynactis carnea            -                     -
Ectopleura crocea            -                     -
Sertularella                 -             0.1 [+ or -] 0.1
  mediterranea
Plumularia setacea           -                     -
Tripalea clavaria            -                     -
Mytilus platensis    54.5 [+ or -] 5.8     43.3 [+ or -] 5.7
Arbacia dufresnei     0.1 [+ or -] 0.05    0.03 [+ or -] 0.03
Asterina sterifer     0.1 [+ or -] 0.06     0.3 [+ or -] 0.1

                                  Natural reef

                             BM                    BA

Ulva lactuca                  -                    -
Corallina                     -                    -
  officinalis
Polysiphonia sp.              -                    -
Rhodymenia sp.                -                    -
Porifera sp. 1                -                    -
Porifera sp. 2                -                    -
Porifera sp. 3                -                    -
Porifera sp. 4                -                    -
Porifera sp.5                 -                    -
Porifera sp. 6                -                    -
Amphisbetia            2.4 [+ or -] 0.5     0.5 [+ or -] 0.2
  operculata
Anthothoe              0.2 [+ or -] 0.1    0.03 [+ or -] 0.03
  chilensis
Aulactinia sp.                -                    -
Corynactis carnea             -                    -
Ectopleura crocea             -                    -
Sertularella           0.1 [+ or -] 0.1            -
  mediterranea
Plumularia setacea     1.2 [+ or -] 0.3     2.1 [+ or -] 0.4
Tripalea clavaria     11.7 [+ or -] 1.0    0.57 [+ or -] 0.3
Mytilus platensis     12.1 [+ or -] 2.7    32.7 [+ or -] 4.6
Arbacia dufresnei     0.07 [+ or -] 0.04   0.03 [+ or -] 0.03
Asterina sterifer      0.1 [+ or -] 0.05    0.1 [+ or -] 0.04

                                 Natural reef

                            BAS                  PS

Ulva lactuca                 -                    -
Corallina                    -                    -
  officinalis
Polysiphonia sp.             -                    -
Rhodymenia sp.               -                    -
Porifera sp. 1               -                    -
Porifera sp. 2               -                    -
Porifera sp. 3               -                    -
Porifera sp. 4               -                    -
Porifera sp.5                -                    -
Porifera sp. 6               -                    -
Amphisbetia                  -            0.9 [+ or -] 0.4
  operculata
Anthothoe                    -            0.4 [+ or -] 0.2
  chilensis
Aulactinia sp.               -                    -
Corynactis carnea    0.6 [+ or -] 0.3             -
Ectopleura crocea            -                    -
Sertularella                 -            0.1 [+ or -] 0.07
  mediterranea
Plumularia setacea   0.5 [+ or -] 0.2     0.2 [+ or -] 0.1
Tripalea clavaria    7.9 [+ or -] 0.9    0.03 [+ or -] 0.03
Mytilus platensis            -           32.3 [+ or -] 4.6
Arbacia dufresnei            -           0.07 [+ or -] 0.05
Asterina sterifer    0.1 [+ or -] 0.08    0.1 [+ or -] 0.06

                                Artificial reef

                            JCH                  JCV

Ulva lactuca                 -                    -
Corallina                    -                    -
  officinalis
Polysiphonia sp.             -                    -
Rhodymenia sp.        1.1 [+ or -] 0.2            -
Porifera sp. 1       -                            -
Porifera sp. 2       0.67 [+ or -] 0.15           -
Porifera sp. 3        0.4 [+ or -] 0.1            -
Porifera sp. 4               -             4.4 [+ or -] 1.0
Porifera sp.5                -                    -
Porifera sp. 6               -                    -
Amphisbetia           1.9 [+ or -] 0.4            -
  operculata
Anthothoe             1.5 [+ or -] 0.3    34.1 [+ or -] 7.6
  chilensis
Aulactinia sp.               -                    -
Corynactis carnea            -                    -
Ectopleura crocea            -             1.2 [+ or -] 0.3
Sertularella           08 [+ or -] 0.2            -
  mediterranea
Plumularia setacea    1.0 [+ or -] 0.2            -
Tripalea clavaria            -                    -
Mytilus platensis    36.1 [+ or -] 8.0     2.4 [+ or -] 0.5
Arbacia dufresnei     0.2 [+ or -] 0.05           -
Asterina sterifer     0.4 [+ or -] 0.08           -

                     Artificial reef

                             CR

Ulva lactuca                 -
Corallina                    -
  officinalis
Polysiphonia sp.             -
Rhodymenia sp.               -
Porifera sp. 1               -
Porifera sp. 2               -
Porifera sp. 3               -
Porifera sp. 4               -
Porifera sp.5         0.3 [+ or -] 0.3
Porifera sp. 6       0.17 [+ or -] 0.1
Amphisbetia           3.1 [+ or -] 1.1
  operculata
Anthothoe             3.1 [+ or -] 2.7
  chilensis
Aulactinia sp.               -
Corynactis carnea            -
Ectopleura crocea    0.17 [+ or -] 0.16
Sertularella                 -
  mediterranea
Plumularia setacea   0.13 [+ or -] 0.1
Tripalea clavaria     3.0 [+ or -] 1.0
Mytilus platensis            -
Arbacia dufresnei    0.18 [+ or -] 0.03
Asterina sterifer    0.03 [+ or -] 0.03

Table 4. R values of the ANOSIM test (global r = 0.514, P < 0.1%)
for every pair comparison among sites (natural and artificial
reefs) according to its biological composition. Significant values
in bold (P < 0.1%). Natural outcrops: Restingas Faro (R0), Primera
(R1), Segunda (R2) and Tercera (R3), Banco Pescadores Sur (PS),
Banco del Medio (BM), Banco de Afuera (BA) and the sedimentary slope
in BA (BAS). Artificial reefs: James Clunies horizontal (JCH) and
vertical (JCV), and Cristo Rey (CR).

Tabla 4. R valores del test ANOSIM (global r = 0,514, P < 0,1%)
para cada comparacion pareada entre sitios (arrecifes naturales y
artificiales) acorde a su composicion biologica. Valores significativos
en negrita (P < 0,1%). Arrecifes naturales: Restingas Faro (R0),
Primera (R1), Segunda (R2) y Tercera (R3), Banco Pescadores Sur (PS),
Banco del Medio (BM), Banco de Afuera (BA) y elevacion de roca
sedimentaria en BA (BAS). Arrecifes artificiales (pecios): James Clunies
horizontal (JCH) y vertical (JCV), y Cristo Rey (CR).

       R0     R1      R2      R3     PS     BM    BA     BAS    JCH
R0
R1    0,39
R2    0,76   0,29
R3    0,66   0,19   -0,005
PS    0,70   0,19    0,05    0,07
BM    0,85   0,47    0,68    0,67   0,55
BA    0,72   0,21    0,08    0,13   0,01   0,57
BAS   0,82   0,58    0,94    0,86   0,88   0,33   0,87
JCH   0,50   0,02    0,24    0,19   0,20   0,58   0,23   0,72
JCV   0,86   0,47    0,99    0,88   0,94   0,99   0,98   0,91   0,69
CR    0,55   0,29    0,76    0,58   0,64   0,38   0,62   0,29   0,28

       JCV
R0
R1
R2
R3
PS
BM
BA
BAS
JCH
JCV
CR    0,56

Table 5. SIMPER("similarity percentages" analysis) results for the
natural and artificial reefs of the study area(average similarity
within groups in brackets). Typical species(those contributing more
than 90% to the average similarity for each group) are listed (%
contribution in brackets), Av. Abund.= average cover data, Contrib. %
= contribution in percentage, Cum. % = cumulative percentage.

Tabla 5. Resultados del SIMPER("similarity percentages" analysis)
para los arrecifes naturales y artificiales del area de estudio
(similitud promedio entre grupos entre parentesis) y lista de las
especies tipicas (aquellas que contribuyen en mas de un 90% de la
similitud promedio para cada grupo). Av. Abund.= promedio datos de
cobertura, Contrib. % = contribucion en porcentaje, cum. % =
porcentaje acumulativo.

Substrate type     Species                  Av. Abund.   Contrib. %

Natural (28.01%)   Mytilus platensis        29.49        83.65
                   Tripalea clavaria        3.14         12.31

Artificial

JCH (23.56%)       Mytilus platensis        36.67        96.18
JCV (70.23%)       Anthothoe chilensis      72.37        99.34
CR (18.6%)         Tripalea clavaria        4.40         57.58
                   Amphisbetia operculata   4.55         40.32

Substrate type     Species                  Cum. %

Natural (28.01%)   Mytilus platensis        83.65
                   Tripalea clavaria        95.95

Artificial

JCH (23.56%)       Mytilus platensis        96.18
JCV (70.23%)       Anthothoe chilensis      99.34
CR (18.6%)         Tripalea clavaria        57.58
                   Amphisbetia operculata   97.90

Table 6. Relative abundance (Ra) and frequency (F) of fishes from
natural reefs (Restigas y Bancos) and artificial reefs in the study
area. Ra code: occasional (o) = a single individual, scarce (s) =
2-5 individuals, abundant (a) = 6-10 and very abundant (va) = more
than 10 individuals observed during the immersion. F code: less
frequent (lf) = recorded in less than 20 %; frequent (f) = recorded
between 20-50 % and very frequent (vf) = recorded in more of the 50
% of total dives in each place.

Tabla 6. Abundancia relative (Ra) y frecuencia (F) de los peces de los
arrecifes naturales (restigas y bancos) y los arrecifes artificiales
del area de estudio. Codigo de Ra: ocasional (o): un individuo, escasos
(s): 2-5 individuos, abundantes (a): 6-10 y muy abundantes (va): mas de
10 individuos observados durante la inmersion. Codigo de F: poco
frecuente (lf): registrada en menos del 20%; frecuente (f): registrada
entre el 20-50% y muy frecuente (vf): registrada en mas del 50% de
total de inmersiones en cada lugar.

                                         Natural reef

Species            Common name    Restingas       Bancos    J. Clunies

                                     Ra   (F)    Ra   (F)

Acanthistius       Argentine         s    (f)    a    (f)       va
  brasilianus        sea bass
Bovichthys                           o    (lf)   --             --
  argentinus
Conger                               --          o    (lf)      --
  orbignyamus
Diplodocus         White bream       Va   (vf)   s    (vf)      va
  argenteus
Discopyge          Electric ray      --          o    (lf)      --
  tschudii
Dules auriga                         s    (vf)   s    (f)       s
Pagrus pagrus      Red porgy         s    (vf)   a    (vf)      s
Paralichthys       Flounder          o    (lf)   o    (lf)      --
  patagonicus
Pinguipes          Brazilian         Va   (vf)   va   (vf)      va
  brasilianus      sea bass
Pseudopercis       Brazilian         --          0    (lf)      s
  semifasciata     sandperch
Rajiidae           Eagle ray         o    (lf)   o    (lf)      o
  species indet.
Seriola lalandi    Yellowtail        --          ma   (lf)      --
Squatina sp.       amberjack         --          o    (lf)      --
Triathalassothia                     --          o    (lf)      --
  argentina
Urophycis          Brazilian         --          o    (lf)      o
  brasiliensis     codling
Number of                            10          16             5
  census

                        Artificial reef

Species              J. Clunies   Cristo Rey

                                   Ra   (F)

Acanthistius            (vf)       va   (vf)
  brasilianus
Bovichthys                         --
  argentinus
Conger                             s    (f)
  orbignyamus
Diplodocus              (vf)       va   (vf)
  argenteus
Discopyge                          --
  tschudii
Dules auriga            (lf)       s    (lf)
Pagrus pagrus           (lf)       s    (lf)
Paralichthys                       --
  patagonicus
Pinguipes               (vf)       va   (vf)
  brasilianus
Pseudopercis             (f)       s    (f)
  semifasciata
Rajiidae                (lf)       o    (lf)
  species indet.
Seriola lalandi                    --
Squatina sp.                       --
Triathalassothia                   --
  argentina
Urophycis               (lf)       o    (lf)
  brasiliensis
Number of                          4
  census
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Title Annotation:texto en ingles
Author:Genzano, Gabriel; Giberto, Diego; Bremec, Claudia
Publication:Latin American Journal of Aquatic Research
Date:Nov 1, 2011
Words:7444
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