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And Pomatoschistus Minutus (Pallas, 1770) in a Temperate Estuary, Portugal

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And Pomatoschistus Minutus (Pallas, 1770) in a Temperate Estuary, Portugal Estuarine, Coastal and Shelf Science 66 (2006) 231e239 www.elsevier.com/locate/ecss Feeding ecology, population structure and distribution of Pomatoschistus microps (Krøyer, 1838) and Pomatoschistus minutus (Pallas, 1770) in a temperate estuary, Portugal R. Leita˜o a,*, F. Martinho a, J.M. Neto a, H. Cabral b, J.C. Marques a, M.A. Pardal a a Institute of Marine Research (IMAR), c/o Department of Zoology, University of Coimbra, 3004-517 Coimbra, Portugal b Instituto de Oceanografia, Faculdade de Cieˆncias da Universidade de Lisboa, Campo Grande, 1749-016 Lisboa, Portugal Received 25 April 2005; accepted 30 August 2005 Available online 23 November 2005 Abstract The feeding ecology, growth and spatialetemporal abundance and distribution of Pomatoschistus microps (Krøyer, 1838) and Pomatoschistus minutus (Pallas, 1770) were studied between June 2003 and June 2004, based on beam trawl surveys and macrobenthic samples conducted in the Mondego estuary, Portugal. Polychaetes, molluscs and amphipods were the most important items in the diet of P. microps, while for P. minutus the dominant preys were polychaetes, mysids and decapods. Pomatoschistus microps’ recruitment lasted for six months and was composed of three new cohorts per year. Pomatoschistus minutus had an uncommon long reproductive season, from April to November, and population seg- regation was found, corresponding to the two reproductive peaks. Populations of both species were composed mainly of 0-group individuals with sand gobies presenting a more extended life span. In the Mondego estuary, inter- and intra-specific spatial segregation occurred between the two species and between the 0C and 1C age groups of P. microps. Ó 2005 Elsevier Ltd. All rights reserved. Keywords: Pomatoschistus; diet overlap; life cycle; competition; resource partitioning; Mondego estuary 1. Introduction this resource. The study of diet is often used to evaluate com- petition between fish species (Elliott et al., 2002). High fish abundance concentrated in particular areas, is Trophic and habitat segregation are among the most impor- a common feature in estuarine fish communities (Elliott tant aspects of resource partitioning in fish communities but it et al., 2002). Consequently, co-occurrence of several similar has been difficult to assess and prove the main force behind morphological types (e.g. species of Soleidae, Mugilidae, Go- this process (Ross, 1986a). Evidences of coexistent Gobiidae biidae) with overlap of ecological niches within the communi- species exhibiting vertical spatial segregation, temporal segre- ties is expected. Spatial and temporal distribution, abundance gation in spawning and absence of niche shifts when one patterns of fish assemblages as well as resource partitioning, species was temporarily absent have been reported by Norte- are therefore, important and interesting issues for understand- Campos and Temming (1994). The Gobiidae is a particularly ing the structure and dynamics of these communities (Cabral, successful family in both tropical and temperate seas, well rep- 2000). Competition exists when the requirements of two or resented in estuarine and freshwater environments. Gobiidae more individuals for a particular resource exceed the supply present more than 2000 species (Bouchereau and Guelorget, of that resource in the environment; or, if their demands do 1998) and virtually all gobies exhibit demersal eggs, male not exceed the supply, but they compete directly to obtain parental care and highly variable number of spawnings per season (Miller, 1984). * Corresponding author. In the Mondego estuary, four species of Gobiidae are pres- E-mail address: [email protected] (R. Leita˜o). ent: Pomatoschistus microps (Krøyer, 1838), Pomatoschistus 0272-7714/$ - see front matter Ó 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.ecss.2005.08.012 232 R. Leita˜o et al. / Estuarine, Coastal and Shelf Science 66 (2006) 231e239 minutus (Pallas, 1770), Gobius niger Linnaeus, 1758, and Aphia estuary, with an area of 3.4 km2, and a Mediterranean temper- minuta (Risso, 1810), the two Pomatoschistus species being the ate climate. The estuary comprises two arms, north and south, most abundant fishes in the Mondego estuary assemblage with very different hydrologic characteristics suffering also (Leita˜o et al., in press). The common goby, P. microps, is a sed- different human impacts. The northern arm is deeper, 5e entary species of semi-enclosed lagoon-like environments, 10 m deep at high tide (tidal range 1e3 m), highly hydrody- which undertakes thermic migration in northern Europe (Jones namic and constitutes the main navigation channel and the lo- and Miller, 1966). Although it is usually an annual species in the cation of the Figueira da Foz harbour. The southern arm is Mediterranean Sea (Bouchereau and Guelorget, 1998; Pampou- shallower (2e4 m during high tide, tidal range 1e3 m), and lie, 2001), it can live up to two years in the northern Europe is characterized by large areas of intertidal flats exposed dur- (Jones and Miller, 1966). The main food items for this fish con- ing low tide. sist of meiofauna and small macrofauna (Pihl, 1985; Salgado Fish were collected, monthly, between June 2003 and June et al., 2004). Similarly, the sand goby, P. minutus, is a ubiquitous 2004, using a 2 m beam trawl, with one tickler chain and estuarine and inshore marine fish with a short generation time 5 mm stretched mesh size in the cod end. Sampling was car- (12e18 months) being able to spend its entire life cycle within ried out during the night, at low water of spring tides, in an estuary (Healey, 1971). In both species, young gobies start five stations (A, B, C, D and E) (Fig. 1). Each survey consisted their life form as suprabenthic organisms, changing to epi- of three hauls, at each of the five sampling stations, in a total benthic with one or two months old (Zander, 1990). of 15 min per station. All fish captured were identified, mea- Considering the importance of these species in the marine sured (total length) and weighted (wet weight). Stomachs and estuarine ecosystems (Miller, 1972), especially their role were removed and contents preserved in 4% buffered formalin as intermediate predators in the food-web connecting micro- for later identification. Each prey item was identified to the benthos with large predator fish (Raffaelli and Milne, 1987; lowest taxonomic level possible, counted and weighted (wet Raffaelli et al., 1989), few studies (Arruda et al., 1993; weight). Temperature, salinity, pH and dissolved oxygen Salgado et al., 2004) have focused on the ecology of these spe- near the bottom were measured during sampling. Algae col- cies in the Atlantic coast of Southern Europe (the literature is lected together with fish in beam trawl samples were also particularly scarce for Pomatoschistus microps). weighted (total wet weight). Each season (summer, autumn, The purpose of the present study was (a) to analyze the dis- winter and spring), sediment samples were collected using tribution and abundance patterns of Pomatoschistus microps a Van Veen grab in all sampling stations, in order to determine and Pomatoschistus minutus in a temperate estuary; (b) to granulometry and benthic macroinvertebrates biomass (g mÿ2 compare the feeding ecology and growth of P. microps and AFDW e ash free dry weight). Granulometry was determined P. minutus; and (c) to evaluate the potential for competition after incinerating dried sediment samples at 450 C for 7 h, between the two species. being afterwards sorted in a sieve series and weighed accord- ing to grain size. 2. Material and methods 2.1. Study site and sampling procedures 2.2. Feeding The Mondego estuary is located on the Atlantic coast of The stomach contents of 103 Pomatoschistus microps and Portugal (40 08#N, 8 50#W) (Fig. 1). It is a small intertidal 101 Pomatoschistus minutus were analyzed. The relative Fig. 1. Location of sampling stations within the Mondego estuary. R. Leita˜o et al. / Estuarine, Coastal and Shelf Science 66 (2006) 231e239 233 importance of each prey item in the diet was expressed by of this software is based on the probability paper method, as three indices: the numerical index (NI) e percentage of the proposed by Cassie (1963). number of individuals of a prey over the total number of indi- viduals of all preys; the occurrence index (OI) e percentage of non-empty stomachs in which a prey occurred; and the gravi- 2.4. Distribution and abundance patterns metric index (GI) e percentage in wet weight of a prey in re- lation to the total wet weight of all prey. Feeding activity was A canonical correspondence analysis was performed to evaluated by the vacuity index defined as the percentage of evaluate spatial and temporal variation of both Pomatoschistus empty stomachs (Hyslop, 1980). A mixed method, proposed species, using CANOCO 4.0 (Ter Braak, 1988). This multivar- by Hureau (1970) and adapted by Salgado et al. (2004), the di- iate technique allows one to directly relate the pattern of com- etary coefficient (QI) was also used. Defined as QI Z munity variation with the pattern of environmental variation NI ! GI, this method considers both the weight and the num- (Ter Braak and Prentice, 1988). According to the some au- ber of ingested prey, classifying them into dominant thors, it has several advantages when compared with other di- (QI R 200), secondary (200 O QI R 20) or unimportant rect gradient analysis techniques, in particular when species (QI ! 20). have nonlinear and unimodal relationships to environmental Diet overlap was measured using the Schoener index (SI): gradients. Each species abundance data was averaged by age group (0C and 1C), by sampling area (stations A, B, C, D ! Xn and E) and seasons (summer, autumn, winter and spring). Z SI 1 ÿ 0:5 piA ÿ piB Temperature, salinity, dissolved oxygen, sediment type, algae iZ1 cover, depth at high tide and benthic macroinvertebrates bio- mass were included in the analysis as environmental data.
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