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Sizefrequency Distributions of Joania Cordata and Argyrotheca Cuneata

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Sizefrequency Distributions of Joania Cordata and Argyrotheca Cuneata Marine Ecology. ISSN 0173-9565 ORIGINAL ARTICLE Size-frequency distributions of Joania cordata and Argyrotheca cuneata (Brachiopoda: Megathyrididae) from the Central Tyrrhenian Sea Francesca Evangelisti, Paolo G. Albano & Bruno Sabelli Department of Biological, Geological and Environmental Sciences, University of Bologna, Bologna, Italy Keywords Abstract Argyrotheca cuneata; Brachiopoda; Joania cordata; Recent brachiopods; Secche di Tor Size-frequency distributions can support reliable inferences concerning popula- Paterno; size-frequency distributions. tion dynamics of brachiopods, but only a few data are available so far. In this study, length and width frequency distributions of dead specimens of the Correspondence Recent brachiopods Joania cordata and Argyrotheca cuneata from the Marine Francesca Evangelisti, Department of Protected Area ‘Secche di Tor Paterno’, Central Tyrrhenian Sea, Italy Biological, Geological and Environmental (41°35′ N, 12°20′ E), are reported in order to add new data about size- Sciences, University of Bologna, Via Selmi 3, frequency distributions of brachiopods. The studied specimens came from 40126, Bologna, Italy. E-mail: [email protected] death assemblages in the coralligenous substrate, in the Posidonia oceanica meadows, and in the sand channels. The observed patterns vary from left- Accepted: 11 July 2013 skewed (J. cordata) to right-skewed (A. cuneata), indicating respectively a low and high mortality of smaller individuals. Significant differences between the doi: 10.1111/maec.12096 coralligenous substrate and the P. oceanica meadow were observed for both species, revealing a variation among different habitats. All length and width distributions are clearly polymodal, but the biological meaning of the peaks is difficult to interpret, as the two species seem to have a 2-year life span. A biometric analysis of shell sizes revealed that length and width are the most variable parameters during the growth of the animal. Bambach 1975; Cadee 1982) and taphonomic processes Introduction (Emig 1990; Simoes~ et al. 2005; Tomasovych & Rothfus Size-frequency distributions (SFDs) have been largely 2005). Also, predation can influence SFDs of brachiopods used as a tool in the study of population dynamics of as predators often choose their prey by size, and preda- both fossil (e.g. Cadee 1982; Alvarez 1986, 1990; Bitner & tion pressure on both recent and fossil brachiopods can Pisera 2000; Bitner 2002; Tomasovych 2004) and present- be appreciable (e.g. Evangelisti et al. 2011; Harper 2011). day brachiopods (e.g. Thayer 1975, 1977; Noble & Logan However, growth and mortality rates are the main prob- 1981; Curry 1982; Taddei Ruggiero 1987; Brey et al. lems because their assessment depends on age determina- 1995). Published works showed that distributions patterns tion and there is no convenient way to estimate the age are mainly bi- or polymodal and that the most common of brachiopods. A direct relationship between size and type of population size-structure is a right-skewed one age cannot be the rule because individuals of the same (e.g. Taddei Ruggiero 1987; Bitner 2002), with only a few age may reach different sizes (Thayer 1977), and there cases of left-skewed SFDs (Thayer 1975; Brey et al. 1995). may also be size variations be due to different local envi- Nevertheless, inferences about population dynamics of ronmental conditions, geographic locations or habitats brachiopods by SFDs may provide ambiguous interpre- (e.g. Thayer 1977; Aldridge 1981). Variations in the size tations as size-frequency graphs can be biased by several of living brachiopods have been recently observed factors, such as inadequate sampling (Richards & with latitude and depth (Peck & Harper 2010). Another Marine Ecology 35 (2014) 377–386 ª 2013 Blackwell Verlag GmbH 377 Size-frequency distributions of J. cordata and A. cuneata Evangelisti, Albano & Sabelli possibility is the estimation of age estimation from bands located 12 km off the coast of Lazio, Italy (41°35′ N, on the shell surface. In a study about population dynam- 12°20′ E), and it covers a total surface of 27 hectares. ics of a dense assemblage of Magellania fragilis from the The top of the reef is at –18 m, and the reef lies on the shelf of the Lazarev sea, Antarctica (Brey et al. 1995), the surrounding mud bottom at a depth of ~70 m. The area bands on the shell surface were interpreted as annual is characterized by two primary habitats typical of the growth marks and treated as size-at-age data. Following Mediterranean Sea: the coralligenous substrate composed this method, other authors published works on living of the accumulation of encrusting algae, and small brachiopods (Peck et al. 1997; Park et al. 2000). Never- meadows formed by the endemic Mediterranean seagrass theless, there is no tangible proof that the bands visible Posidonia oceanica. on the shell surface have been formed seasonally and, Several samples were collected at depths of 20–28 m by moreover, extra, non-annual growth lines can be pro- SCUBA divers through airlift suction samplers, which duced, at least in some species, as reported for Terebratal- were used on the coralligenous substrate, on the rhizome ia transversa (Paine 1969). The best way to define age layer of P. oceanica and in the sand channels, providing would be by recurring observations of the same popula- specimens from the three different death assemblages. tions in their life environment over a period of several The samples were sieved with meshes down to 1 mm, years, used in the techniques of staining and radiography and all retrieved brachiopods were identified, counted in corals, but difficulties in sampling the same site, as and measured using an ocular micrometer. All three well as potential modifications due to human intrusions, dimensions of the shell were recorded as defined by have to be taken into account as possible limitations Williams & Brunton (1997), so that length and width (Tunnicliffe & Wilson 1988; Asgaard & Bromley 1990; were those of the ventral valve, which is usually the Taddei Ruggiero 2001). In some cases, analyses were per- largest one. formed on populations kept alive in the laboratory (e.g. A biometric analysis was carried out on the two species Thayer 1975; Curry 1982; Collins 1991) but the results to investigate their variability, and all three dimensions of can be inaccurate, as reproducing natural life conditions the shell were taken into account. Data were put in artificially is difficult or often impossible. length-width and thickness-width scatter diagrams, which In this study we have investigated the SFDs of two were performed on each kind of sample separately. Recent brachiopods, Joania cordata and Argyrotheca cune- Spearman rank correlations and linear regressions were ata, in order to add new data about SFDs of brachiopods. computed for each scatter diagram to determine the rela- The two species are micromorphic brachiopods living in tionship among the variables. For each species, an analy- shallow waters, attached to cave walls and roofs, boulders sis of covariance (ANCOVA) was also performed to and coralligenous substrate (Logan 1979; Logan et al. highlight possible differences in length-width and thick- 2004; Alvarez et al. 2005, 2008). Their morphology is ness-width relationship among the three analysed sam- illustrated in these publications. The analysis was carried ples. All statistical analyses were performed using the out on samples from the Marine Protected Area ‘Secche program R (R Development Core Team 2008). di Tor Paterno’, Central Tyrrhenian Sea, as part of the Length and width frequency distributions were also biodiversity study of the area (Albano & Sabelli 2011; performed to analyse their variation within and between Evangelisti et al. 2011, 2012). The distributions were the two species. Thickness was not considered because no examined for the two species separately and, within each significant variation among specimens was observed for species, for the three different death assemblages which either species. Histograms were performed for both spe- the specimens came from: within the coralligenous sub- cies, for each kind of sample separately. For each distri- strate, within a Posidonia oceanica meadow, and from bution, a Shapiro–Wilks test and Skewness index were sand channels containing specimens originated from the computed to investigate shape. For each species, length nearby coralligenous substrate and Posidonia meadows. and width frequency distributions of the three samples The outcome was then compared with data from the lit- were compared in their shape and mean values using erature. This is the first study on size-frequency distribu- respectively the Kolmogorov–Smirnov (D) and Mann– tions on Recent specimens of J. cordata and A. cuneata. Whitney (U) tests, to highlight possible differences among the death assemblages of the coralligenous sub- strate, the P. oceanica meadow and the death assemblages Material and Methods in the sand channels. Both tests were computed with the Samples of empty shells of Joania cordata and Argyrotheca program PAST (Hammer et al. 2001). A level of 5% was cuneata were collected from the Marine Protected Area considered significant for all statistical analyses. The stud- ‘Secche di Tor Paterno’, Central Tyrrhenian Sea, from ied material is kept in the Zoological Museum of the May to June 2007. The area is a completely off-shore reef University of Bologna, Italy. 378 Marine Ecology 35 (2014) 377–386 ª 2013 Blackwell Verlag GmbH Evangelisti,
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