Modelling sponge species diversity using a morphological predictor: a tropical test of a temperate model

The contribution of sponges to marine surveys is often underestimated due to problems of identification, synonymous species and limited numbers of specialists in the field. Bell & Barnes (2001) illustrated how sponge morphological diversity (diversity of body forms) might be used as a predictor of sponge species diversity and richness. This study investigated these relationships at six tropical West Indian Ocean localities in a number of habitat types. These habitats included tropical coral reefs, soft substratum (seagrass, mangrove and sand), caves and boulders. Sampling was undertaken at three depth zones in coral reef habitats only (intertidal, 10–15 m and 20–25 m), with the other habitats sampled in less than 10m of water. Species diversity and richness were significantly correlated (P < 0.05) with morphological diversity at all localities and depths in coral reef and soft substratum habitats. However, no significant correlation was found between these variables in cave or boulder habitats. The slope of the linear regression found between morphological diversity and species diversity did not significantly differ between coral reef, soft substratum and temperate reef (data taken from Bell & Barnes 2001) habitats. Similarly coral reefs showed the same relationship between morphological diversity and species richness as temperate reefs, however the relationship between morphological diversity and species richness was significantly different at both habitats compared with soft substratum environments. Sponge morphological diversity therefore may be more useful as a predictor of sponge species diversity, rather than species richness, as the former relationship is common between more habitats than the latter.     

Substratum as a structuring influence on assemblages of Arctic bryozoans

The nature of the substratum is a fundamental factor determining the types of organisms and communities found in many terrestrial and benthic habitats. The extent to which this is true in extreme environments was investigated using bryozoan assemblages as model organisms in an Arctic fjord (Kongsfjorden 79°N, 12°E) in summer 2001 using SCUBA. Twenty-seven substrate samples of 0.25 m² were taken at 10 m depth from the inner glacial basin to the mouth of the fjord. Multivariate analyses revealed four different bryozoan assemblages. The sea floor of the inner basin of Kongsfjorden near the glacial fronts was characterized by low diversity and dominance of the ctenostome species Alcyonidium disciforme Smitt. Highest richness and diversity occurred on rock substratum with mean size >10 cm², on which the most common species was the pioneer Harmeria scutulata Busk (abundance: 15%). On smaller rocks with mean size <2 cm², the runner-like pioneer species Electra arctica Borg comprised most individuals of the assemblage (98%). Yet another pioneer, Celleporella hyalina Linnaeus, was the most abundant species (49%) on substratum dominated by algae. Thus, in each habitat type, pioneers dominated but different species and to different extents. There was much variation in species composition and abundance within assemblages of heterogeneous habitats, and this study emphasizes the importance of microhabitats and physical conditions. Heterogeneity was evident at scales of <1 m.     

Environmental distribution of colony growth form in the favositid Pleurodictyum americanum

Pandolfi, John M. & Burke, Collette D. 1989 01 15: Environmental distribution of colony growth form in the favositid Pleurodictyum americanum. Lethaia , Vol. 22, pp. 69–84. Oslo. ISSN 0024–1164.
Colony growth form is of fundamental importance to understanding the ecology of both modern and ancient marine sessile colonial animals. Fourier shape analysis of the coral Pleurodictyun americantum (Tabulata: Favositida) from the Middle Devonian Hamilton Group of New York State, indicates that colony growth form is variable between environments. Discriminate function analysis of harmonics 2–6 of 51 assemblages of Pleurodictyum americanum (N = 1900) shows that this species displays an onshore to offshore gradient in colony shape. Offshore environments characterized by low levels of turbidity, oxygen, and light contain more flattened, less hemispherical growth forms, whereas onshore environments characterized by high levels of turbidity, oxygen, and light contain more hemispherical, less flattened growth forms. Harmonic shape analysis detected subtle differences among samples of P. americanum from different environments, but also showed that distinctive morphotypes are distributed within horizons. as well as between them. In fact, no one-to-one correspondence in growth form to environment is apparent; growth forms are distributed within environments, suggesting that genetic factors may have had a greater influence over coral growth form than environment. In tabulate corals, patterns of within species variability must be determined before growth form may be useful in interpreting ancient environments. □ Tabulate coral, Fourier shape analysis, morphological variability, growth form, Devonian, Hamilton Group, New York .     

Relationships between coral reef substrata and fish

 The objective of this work is to identify which substrata characteristics (such as coral morphology, coral diversity, coral species richness, percentage coverage by live coral or by algae) influence the structure and abundance of fish communities. The study was carried out at Reunion Island, Indian Ocean, where six sites were sampled in two zones (reef flat and outer reef slope). Quantitative data were collected by visual census techniques, along a linear transect of 50 m for the substratum, and a belt of 50×2 m for the fish communities. Correspondence analysis (CA) and an optimising cluster analysis, called dynamic clustering method (DCM) were used to describe and compare fish assemblages with the benthic composition. The relationships between benthic and fish communities were examined using the classes revealed by the partitioning of the substratum with DCM. This partitioning allowed us to derive four classes of substratum: the non-disturbed reef flat, the non-disturbed outer reef slope, the perturbed reef habitat and the reef pass. The analysis of the partitioning based on the coral variables suggests that there are significant relationships between benthic and fish assemblages. Accepted: 26 July 1996     

Changes in Coral Reef Communities and an Associated Reef Fish Species, <Emphasis Type="Italic">Cephalopholis cruentata</Emphasis> (Lacépède), After 30 years on Curaçao (Netherlands Antilles)

Using the same methodology and identical sites, we repeat a study dating from 1973 and quantify cover of hard coral species, soft corals, sponges, hard substratum and soft substratum, and density of a commercially important reef fish species, the graysby Cephalopholis cruentata, along a depth-gradient of 3–36 m on the coral reefs of Curaçao. The objective was to determine the multi-decade change in benthic coral reef cover and structural complexity, and their effect on densities of an associated reef fish species. Total hard coral cover decreased on average from 52% in 1973 to 22% in 2003, representing a relative decline of 58%. During this time span, the cover of hard substratum increased considerably (from 11 to 58%), as did that of soft corals (from 0.1 to 2.2%), whereas the cover of sponges showed no significant change. Relative decline of hard coral cover and of reef complexity was greatest in shallow waters (near the coast), which is indicative of a combination of anthropogenic influences from shore and recent storm damage. Cover of main reef builder coral species (Agaricia spp., Siderastrea siderea, Montastrea annularis) decreased more than that of other species, and resulted in a significant decrease in reef complexity. Although density of C. cruentata was highly correlated to cover of Montastrea and Agaricia in 1973, the loss of coral cover did not show any effect on the total density of C. cruentata in 2003. However, C. cruentata showed a clear shift in density distribution from shallow water in 1973 to deep water in 2003. It can be concluded that the reefs of Curaçao have degraded considerably in the last three decades, but that this has had no major effect on the population size of one commercially important coral-associated fish species.     

A coral-bryozoan association from the Neogene of northwestern Europe

The Neogene cheilostomatous bryozoan Celleporaria palmata lived facultatively in association with the scleractinian coral Culicia. For Culicia the association was obligatory; the symbiosis was most probably mutualistic. Benefits for the coral appear to have been: (1) the bryozoan provided substrate in a soft-bottom environment; (2) the coral could feed at higher levels above the sediment; (3) the feeding currents generated by the bryozoan provided the corals with food (no competition for food would occur as the coral would collect larger prey); and (4) the surrounding of the cordites by the bryozoan provided strength and lateral protection. Benefits for the bryozoan appear to have been: (1) the stinging cells of the coral provided protection against predators and settlers; (2) the positioning of the corals on the maculae saved the bryozoan subsequent energy expenditure in further construction of polymorphs and secondary calcification. Negative effects for the bryozoan might have been the inhibition of possible functions of the maculae other than as exhalant chimneys, e.g., production and generation of male gametes. *** Symbiosis, mutualism, C ulicia , C ryptangia , C elleporaria , Bryozoa, Scleractinia.     

Bryozoaires des milieux récifaux miocèes du sillon sud-rifain au Maroc

Moissette, P. & Saint Martin, J_:P. 1995 11 30 Bryozoaires des milieux récifaux miocenes du sillon sud-rifain au Maroc. [Bryozoans from Miocene reef environments in the South-Rifian corridor of Morocco. In late Miocene coral reefs from the Fes area (Northern Morocco), bryozoans are the best-represented group. An inventory of the bryozoan fauna (59 species) was taken on two reefs with different organization and palaeogeographical situation in a late Miocene seaway between the Atlantic and the Mediterranean. A study of the bryozoans and their zoarial forms allows an improved reconstruction of reefal environments and available habitats. Membraniporiforrn (encrusting) colonies largely predominate, whereas celleporiforms (nodular colonies) are more rare. Rigid erect (vinculariiform, adeoniform, and reteporiform) and rigid articulated zoarial types (cellariiform and catenicelliform) are fairly well represented. At depths of about 10 m and on the reef front encrusting bryozoans were well developed in cryptic habitats on living coral substrates. Erect species lived in cavities provided by the coral framework, and plants offered flexible substrates for the attachment of epiphytal forms. □Bryozoa, coral reefs, Morocco, Messininn, pulueoecology. Dans les récifs coralliens du Miocène supérieur de la région de Fes (Maroc septentrional), les bryozoaires constituent, parrni les organismes conservés, le groupe le mieux représenté. Un recensement de la fame de bryozoaires (59 espèces) a été effectué dans deux récifs d'organisation et de situation paléogéographique différentes, dans un secteur de communication entre Atlantique et Méditerranée au MiocéFne supérieur. L'étude des bryozoaires et de leurs formes zoariales permet de proposer une meilleure reconstitution de I'environnement récifal et des habitats disponibles. Les colonies membraniporiformes (encrobtantes) predominent largement, alors que les cellepori-formes (colonies noduleuses) sont beaucoup plus rares. Les types zoariaux érigés rigides (vincula-riiforme, adéoniforme et rétéporiforme) et érigés articulés (cellariiforme et catenicelliforme) sont assez bien représentés. A des profondeurs denviron 10 m et dans la partie antérieure des constructions récifales, les bryozoaires encroûtants étaient bien développés dans des habitats cryptiques sur substrats coralliens vivants. Des formes dressées vivaient dans les cavités ménagées par le bâti corallien et des supports végétaux permettaient également la fixation de formes épiphytes. □Bryozouires, récifs coralliens, Muroc, Messinien, puléoécologie.     

Bryozoan assemblages in Messinian deposits of western Algeria

Messinian (Uppermost Miocene) outcrops near the town of Oran in western Algeria yielded rich bryozoan faunas: 188 species and forms. Bryozoans were studied in ten different deposits and 27 horizons representing various marly and calcareous facies. The number of species is lowest in the coral reefs (51), intermediate in the coralline-algae limestones (95), and highest in the mark(167). Also, zoarial growth forms are more diverse in marly deposits than in calcareous ones. These differences are interpreted in terms of bottom substrate, depth, deposition rate, and hydrodynamic conditions. An analysis based on zoarial growth forms and known bathymetric ranges of species still living gives more precise depth indications for each studied assemblage. A detailed survey of these assemblages in several sections shows that a gradual lowering of regional sea level occurred during the Messinian. From the presence of a number of warm-water bryozoan taxa, a higher temperature than in the Present Mediterranean sea is also favored. □ Algeria, Bryozoa, calcareous algae, mark Messinian, paleoecology, reefs.     

Fish assemblages of Cais do Carvão Bay (Madeira Island) determined by the visual census technique

Fish assemblages in Cais do Carvão Bay, Madeira Island, a proposed marine protected area (MPA), were determined from a diver visual census. A total of 32 transect counts were performed. Habitats sampled included sandy bottom, rocky boulders, vertical walls and rocky outcrops. Species richness, diversity, density, trophic structure, size and spatial organization were documented for the fish assemblages. Forty‐four species from 23 families were encountered; 32% belonged to Sparidae (10) and Labridae (four). The greatest species richness (25) was observed in rocky boulder habitat at 10–15 m depth, while the lowest (five) occurred over a deeper sand habitat. The greatest density (760·5 individuals per 100 m²) was recorded over rocky outcropping (20–25 m deep), and the lowest of 11·6 individuals per 100 m² was over a sand bottom at 10–15 m depth. Thalassoma pavo , Abudefduf luridus and Chromis limbata had higher densities on hard bottoms, while Heteroconger longissimus was the most abundant species in sand bottom habitats. No significant differences were detected for all indices calculated among depth intervals for sand and rocky boulder stations. Sand and rock boulder substratum, however, differed significantly for the 10–15 m depth stratum.     

IMPORTANCE OF MACRO‐ VERSUS MICROSTRUCTURE IN MODULATING LIGHT LEVELS INSIDE CORAL COLONIES1

Adjusting the light exposure and capture of their symbiotic photosynthetic dinoflagellates (genus Symbiodinium Freud.) is central to the success of reef‐building corals (order Scleractinia) across high spatio‐temporal variation in the light environment of coral reefs. We tested the hypothesis that optical properties of tissues in some coral species can provide light management at the tissue scale comparable to light modulation by colony architecture in other species. We compared within‐tissue scalar irradiance in two coral species from the same light habitat but with contrasting colony growth forms: branching Stylophora pistillata and massive Lobophyllia corymbosa. Scalar irradiance at the level of the symbionts (2 mm into the coral tissues) were <10% of ambient irradiance and nearly identical for the two species, despite substantially different light environments at the tissue surface. In S. pistillata, light attenuation (90% relative to ambient) was observed predominantly at the colony level as a result of branch‐to‐branch self‐shading, while in L. corymbosa, near‐complete light attenuation (97% relative to ambient) was occurring due to tissue optical properties. The latter could be explained partly by differences in photosynthetic pigment content in the symbiont cells and pigmentation in the coral host tissue. Our results demonstrate that different strategies of light modulation at colony, polyp, and cellular levels by contrasting morphologies are equally effective in achieving favorable irradiances at the level of coral photosymbionts.     

Variation in symbiont distribution between closely related coral species over large depth ranges

Symbiotic algae in coral species distributed over a large depth range are confronted with major differences in light conditions. We studied the genetic variation of Symbiodinium in the coral genus Madracis over depth (5-40 m) and at two different colony surface positions. Using polymerase chain reaction-denaturing gradient gel electrophoresis ITS2 nuclear ribosomal DNA analyses, we consistently identified three symbiont genotypes with distributions that reveal patterns of host specificity and depth-based zonation. ITS2 type B7 Symbiodinium is the generalist type, occurring in all zooxanthellate Madracis corals and at all depths. Type B13 is restricted to the shallow water specialist Madracis mirabilis. Type B15 is typical of deep reef environments and replaces B7 in the depth generalist Madracis pharensis. Contrasting with variation over depth, we found strong functional within-colony uniformity in symbiont diversity. Relating symbiont distributions to measured physical factors (irradiance, light spectral distribution, temperature), suggests depth-based ecological function and host specificity for Symbiodinium ITS2 types, even among closely related coral species.     

Tectonic subsidence provides insight into possible coral reef futures under rapid sea-level rise

Sea-level rise will change environmental conditions on coral reef flats, which comprise extensive habitats in shallow tropical seas and support a wealth of ecosystem services. Rapid relative sea-level rise of 0.6 m over a relatively pristine coral reef in Solomon Islands, caused by a subduction earthquake in April 2007, generated a unique opportunity to examine in situ coral reef response to relative sea-level rise of the magnitude (but not the rate) anticipated by 2100. Extent of live coral was measured from satellite imagery in 2003, 2006, 2009 and 2012. Ecological data were obtained from microatolls and ecological surveys in May 2013. The reef was sampled at 12 locations where dense live hard coral remained absent, remained present or changed from absent to present following subsidence. Ecological data (substratum depth, live coral canopy depth, coral canopy height, substratum suitability, recruitment, diversity and Acropora presence) were measured at each location to identify factors associated with coral response to relative sea-level rise. Vertical and horizontal proliferation of coral occurred following subsidence. Lateral expansion of live coral, accomplished primarily by branching Acropora spp., resulted in lower diversity in regions which changed composition from pavement to dense live coral following subsidence. Of the ecological factors measured, biotic factors were more influential than abiotic factors; species identity was the most important factor in determining which regions of the reef responded to rapid sea-level rise. On relatively pristine reef flats under present climatic conditions, rapid relative sea-level rise generated an opportunity for hard coral to proliferate. However, the species assemblage of the existing reef was important in determining response to sea-level change, by providing previously bare substrate with a source of new coral colonies. Degraded reefs with altered species composition and slower coral growth rates may be less able to respond to climate change-induced sea-level changes.     

BRACHIOPOD PALAEOECOLOGY IN MIDDLE DANIAN LIMESTONES AT FAKSE, DENMARK

The Danian limestone in the Fakse quarry displays banks of coral limestone interfingering with banks of bryozoan limestone. (1) Previous and present studies on the brachiopods show that ‘Rhynchonella’ flustracea Buch, ‘R.’ faxensis Posselt, ‘Terebratula’ mobergi Lundgren, and Argyrotheca pindborgi (Nielsen) are restricted to the coral limestone of Fakse. (2) Crania (Danocrania) transversa (Lundgren), C. (D.) tuberculata (Nilsson), Isocrania aff. egnabergensis (Retzius), ‘Terebratula’ fallax Lundgren, and Carneithyris incisa (Buch) are known from bryozoan limestone at Fakse and other Middle Danian localities. (3) Common to both kinds of sediments in Fakse quarry and found at other Middle Danian bryozoan limestone localities as well are Terebratulina aff. chrysalis (Schlotheim), Argyrotheca scabricula (Koenen), Megathiris bruennichi (Rosenkrantz), Platidia sp., and Thecidellina? groenwalli (Nielsen). (4) ‘Terebratula cincta’ Nielsen is restricted to Fakse quarry and occurs in both kinds of sediments. (5) Some of the species restricted to the coral limestone appear to be morphologically adapted to fit in between the branches of the dominant coral in Fakse quarry, Dendrophyllia candelabrum Hennig.     

A Novel Technique for the Reattachment of Large Coral Reef Sponges

Sponges are dominant components of coral reef ecosystems, often exceeding reef-building corals in abundance. Large sponges, often more than 1 m in diameter, may be hundreds to thousands of years old. When damaged or dislodged, large sponges usually die because they are unable to reattach to the reef substratum. Because suitable methods for reattaching dislodged sponges are lacking, they are typically excluded from coral reef restoration efforts. Here we present a novel technique for the reattachment of large sponges that was tested using the Caribbean Giant barrel sponge, Xestospongia muta . Transplants of X. muta were conducted at 15- and 30-m depth off Key Largo, Florida. Despite the active hurricane season of 2005, 90% of deep and 35% of shallow transplants survived, with nearly 80% reattaching to the substratum and growing after 2.3–3 years. This technique may be generally adapted for securing large sponges in coral reef restoration efforts.     

Vertical partitioning of the phytoplankton assemblage in ultraoligotrophic Crater Lake, Oregon, U.S.A.

SUMMARY 1. The summertime phytoplankton assemblage in abysmally deep (Z_max: 589 m) Crater Lake, Oregon, consists of over 100 species, which are variously distributed in the upper 200 m of the vertical water column. The depth distribution of the lake's three most prevalent species follows a predictabk pattern: Nitzschia gracilis in the 0–20 m stratum, Tribonema sp. at mid–depth (80–20 m), and Stephanodiscus hantzschii in the lowermost stratum (160–200 m). These major species, which account for approximately 80% or more of the lake's total phytoplankton biomass and primary production, exist under atypical temperature, light, and nutrient conditions.
2. The spatial distribution of phytoplankton in Crater Lake resembles a three-tier structure. Unlike most lakes, where the entire phytoplankton communities exist in less disparate environmental conditions, or are vertically mixed periodically by storm events and seasonal lake turnover. the Crater Lake community is partitioned into stratified environments.
5. The disparate and unusual characteristics of these environments, and the hydrological and limnological stability of the lake basin, are perhaps important factors regulating the diversity, dominance. and partitioning of the lake's phytoplankton populations.     

Reef fish community structure of the Fernando de Noronha Archipelago (Equatorial Western Atlantic): the influence of exposure and benthic composition

We studied the reef fish assemblage of eight reefs within the oceanic archipelago of Fernando de Noronha, off northeastern Brazil. In a total of 91 belt transects (20 × 2 m) we recorded 60 species from 28 families. The 25 most abundant species accounted for about 98% of all fish recorded in this study and most of these species are widely distributed in the Western Atlantic. The majority of fish counted were planktivores (37.0%), followed by mobile invertebrate feeders (28.5%), territorial herbivores (11.3%), roving herbivores (10.5%), omnivores (7.1%), macrocarnivores (6.5%) and sessile invertebrate feeders (0.03%). In terms of biomass, roving herbivores were the most representative (41.8%), followed by mobile invertebrate feeders (19.9%), macrocarnivores (14.3%), omnivores (14.0%), piscivores (8.3%), planktivores (1.4%), territorial herbivores (0.3%), and sessile invertebrate feeders (0.03%). Overall, density and biomass of fishes were positively correlated with coral cover and depth, and negatively correlated with wave exposure. These relationships are probably a response to the habitat complexity provided by the higher coral cover in deeper reefs (>10 m) of the archipelago or to the lower water turbulence below 10 m deep. Carnivores and mobile invertebrate feeders were mainly influenced by depth and non-consolidated substratum, planktivores and omnivores by wave exposure and herbivores by algal cover. Although our results suggest that habitat characteristics may play a role in determining the distribution of some fish species, we also found several habitat generalists, suggesting that the community is dominated by versatile species.     

Bryozoans, corals, and microfacies of Lower Eifelian (Middle Devonian) limestones at Kierspe, Germany

The Lower Eifelian Meinerzhagener Korallenkalk (= upper Cultrijugatus Beds) at Kierspe, Sauerland, contains a rich reefal fauna. Eight bryozoan species are described, two of them are new: the cystoporate Fistuliporella kierspensis n. sp. and the trepostome Leptotrypella sophiae n. sp. The bryozoans from the Meinerzhagener Korallenkalk shows distinct similarities to the Lower–Middle Devonian of Spain (Santa Lucía Formation, Emsian–Eifelian), and to the Middle Devonian (Eifelian) of Transcaucasia. The coral fauna comprises five tabulate corals and one rugose coral that document a paleobiogeographic relationship between Central and Eastern Europe and Central Asia. The associated fauna is represented by brachiopods, ostracods, and echinoderms. The studied limestones also commonly contain calcimicrobes represented by three species. The faunal and microfacial characteristics indicate a shallow marine depositional environment just above the storm wave base, with a supposed depth of 20 m, within the photic zone. The nutrient regime was at least a mesotrophic. The upper boundary of the Cultrijugatus Beds coincides with the Chote?-Event that strongly affected brachiopods, whereas corals and bryozoans were insensitive to this event.     

The functional importance of <Emphasis Type="Italic">Acropora austera</Emphasis> as nursery areas for juvenile reef fish on South African coral reefs

Many coral reef fish species use mangrove and seagrass beds as nursery areas. However, in certain regions, the absence or scarcity of such habitats suggests that juvenile coral reef fish may be seeking refuge elsewhere. The underlying biogenic substratum of most coral reefs is structurally complex and provides many types of refuge. However, on young or subtropical coral reefs, species may be more reliant on the living coral layer as nursery areas. Such is the case on the high-latitude coral reefs of South Africa where the coral communities consist of a thin veneer of coral overlaying late Pleistocene bedrock. Thus, the morphology of coral species may be a major determinant in the availability of refuge space. Acropora austera is a branching species that forms large patches with high structural complexity. Associated with these patches is a diverse community of fish species, particularly juveniles. Over the past decade, several large (>100 m²) A. austera patches at Sodwana Bay have been diminishing for unknown reasons and there is little evidence of their replacement or regrowth. Seven patches of A. austera (AP) and non-A. austera (NAP) were selected and monitored for 12 months using visual surveys to investigate the importance of AP as refugia and nursery areas. There were significant differences in fish communities between AP and NAP habitats. In total, 110 species were recorded within the patches compared to 101 species outside the patches. Labrids and pomacentrids were the dominant species in the AP habitats, while juvenile scarids, acanthurids, chaetodons and serranids were also abundant. The diversity and abundance of fish species increased significantly with AP size. As the most structurally complex coral species on the reefs, the loss of APs may have significant implications for the recruitment and survival of certain fish species.     

Territorial and non-territorial spawning behaviour in the bream

The spawning behaviour of bream Abramis brama was studied in 1993, in a harbour on the River Meuse, Belgium. Fish spawned from 22 to 27 April and from 11 to 14 May, when the water temperature rose to 14.5) C. The reproductive behaviour of the bream was studied within a 15 m long part of the harbour using a video camera. Territorial males with tubercles (33–43 cm total length; >5 years old) defended bank areas of diameter 80–150 cm which included spawning substratum, i.e. roots of alder and willow trees and aquatic plants. Water depth ranged from 25 to 50 cm. Non-territorial males without tubercles (24–33 cm; 3–4 years old), remained 2–4 m away from the bank. Aggressive behaviour between males was frequent and, occasionally, males with tubercles were unable to defend a territory. Mature females (25–43 cm; >3 years old), coming from the deeper water of the surrounding area, were followed by non-territorial males before spawning in territories near the bank.