A comparison of circatidal rhythmicity and entrainment by hydrostatic pressure cycles in the rock goby, Gobius paganellus L. and the shanny, Lipophrys pholis (L.)

The intertidal teleosts Gobius paganellus and Lipophrys pholis show endogenous circatidal activity rhythms when recorded in constant conditions. Under these conditions, the rhythm of L. pholis is the more precise which may indicate stronger coupling between underlying circalunadian oscillators in this species. In G. pagunellus the inter-oscillator coupling may be weaker and this could enable a more subtle interpretation of tidal fluctuations than in L. pholis . The oscillators may, however, be fundamentally different in the two species; circalunadian in G. paganellus and circatidal in L. pholis .
When exposed to hydrostatic pressure cycles of tidal frequency both species responded pre- dominantly to increasing pressure, which suggests that in the wild they are likely to be most active on the rising tide. Hydrostatic pressure cycles are confirmed as a zeitgeber for both species by the successful entrainment of some individuals. The lack of entrainment of others impIies that additional zeitgebers are required for complete entrainment.     

Juvenile recruitment, growth and maturation of Lipophrys pholis (Pisces: Blenniidae), from the west coast of Portugal

Data on the juvenile recruitment, growth and size at maturation of Lipophrys pholis in Portuguese waters are compared with the information available on the biology of this species at higher latitudes. In Portugal, recruitment extends for a much longer period, young fish grow faster and sexual maturation is earlier than at higher latitudes. There is a delay of 2–3 months between the appearance of the first eggs on the shore and the recruitment of the first juveniles to the pools.     

Phylogenetic relationships of the north-eastern Atlantic and Mediterranean blenniids

The phylogenetic relationships of 27 north-eastern Atlantic and Mediterranean blennioids are analysed based on a total of 1001 bp from a combined fragment of the 12S and 16S mitochondrial rDNA. The most relevant results with implications in current blenniid taxonomy are: (1) Lipophrys pholis and Lipophrys (=  Paralipophrys ) trigloides are included in a well-supported clade that by the rule of precedence must be named Lipophrys ; (2) the sister species of this clade are not the remaining species of the genus Lipophrys but instead a monotypic genus comprising Coryphoblennius galerita ; (3) the smaller species of Lipophrys were recovered in another well-supported and independent clade, which we propose to be recognized as Microlipophrys ; (4) although some authors included the genera Salaria and Lipophrys in a single group we have never recovered such a relationship. Instead, Salaria is more closely related to the genera Scartella and Parablennius ; (5) the genus Parablennius , which was never recovered as a monophyletic clade, is very diverse and may include several distinct lineages; (6) the relative position of Aidablennius sphynx casts some doubts on the currently recognized relationships between the different blenniid tribes. Meristic, morphological, behavioural and ecological characters support our results and are also discussed. The possible roles of the tropical West African coast and the Mediterranean in the diversification of blenniids are discussed.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 283–295.     

Increasing temperature decreases the predatory effect of the intertidal shanny Lipophrys pholis on an amphipod prey

Interactions between Lipophrys pholis and its amphipod prey Echinogammarus marinus were used to investigate the effect of changing water temperatures, comparing current and predicted mean summer temperatures. Contrary to expectations, predator attack rates significantly decreased with increasing temperature. Handling times were significantly longer at 19° C than at 17 and 15° C and the maximum feeding estimate was significantly lower at 19° C than at 17° C. Functional‐response type changed from a destabilizing type II to the more stabilizing type III with a temperature increase to 19° C. This suggests that a temperature increase can mediate refuge for prey at low densities. Predatory pressure by teleosts may be dampened by a large increase in temperature (here from 15 to 19° C), but a short‐term and smaller temperature increase (to 17° C) may increase destabilizing resource consumption due to high maximum feeding rates; this has implications for the stability of important intertidal ecosystems during warming events.     

Interspecific Utility of Microsatellites in Fish: A Case Study of (CT)n and (GT)n Markers in the Shanny Lipophrys pholis (Pisces: Blenniidae) and Their Use in Other Blennioidei

We report the development of new microsatellite markers that can be used for population analyses in the shanny Lipophrys pholis. The procedure involved the construction of a microsatellite-enriched genomic bank. Five (GT)_n and (CT)_n microsatellites have been characterized, four of which are polymorphic. The analysis of one population allowed us to verify their usefulness as markers in population studies. Moreover, interspecific amplifications have been performed using primers defined in other species to amplify Lipophrys pholis, or using the primers defined in Lipophrys pholis to amplify other species. We use these results to discuss the hypothesis that microsatellites are highly conserved in fish. Received May 13, 1999; accepted October 28, 1999.     

Functions of the testicular gland in two blenniid fishes, Salaria (=Blennius) pavo and Lipophrys (= Blennius) dalmatinus (Blenniidae, Teleostei) as revealed by electron microscopy and enzyme histochemistry

The functions of the testicular gland in two different blenniid fishes, Salaria pavo and Lipophrys dulmutinus , are described by fine structural and enzyme histochemical methods. In the testes of the two fishes no mature spermatozoa are found. Sperrniogenesis occurs only until the spermatidal stage. Spermatids are released into the testicular gland. During the spawning period the testicular gland functions in differentiation of spermatids, nutrition of spermatids, and secretion of sialomucins. After spawning, the testicular gland has phagocytotic functions in resorbtion of remaining spermatids, which are transformed into lipids in the gland cells. During the interspawning period the testicular gland is a storage reservoir for lipids and phospholipids which are re-transformed into testicular gland secretion in the next reproductive season. Testicular gland cells themselves do not have steroidogenetic functions, but steroids are synthesized by interstitial cells homologous to Leydig cells in other fish. Possible explanations for the reduction of the testis in L. dalmatinus and implications of the testicular gland in taking over testicular functions are discussed.     

Site fidelity of intertidal fish to rockpools

Gobius paganellus, Lipophrys pholis and Coryphoblennius galerita are wide‐spread intertidal fish that spend their earlier life stages in rock pools, and yet very little is known about their site fidelity behaviour. For these species, fidelity to rockpools may result in increased fitness costs in a predicted scenario of warmer sea water, due to the low thermal inertia of these water bodies. In this context, it is relevant to investigate these species' site fidelity. We made a mark‐recapture study to assess the mentioned species' movements within and between rockpools. We tagged a total of 530 individuals of the aforementioned species with the Visible Implant Elastomer and tracked their movement for a 7‐month period. We found that site fidelity and specific rockpools conditions are important factors in distribution of intertidal blennies and gobies. We also examined the relations between rockpool volume, depth and site fidelity. We found that G. paganellus tends to remain in its original marking pool, with an average recapture rate of 20.5%, but showed no evidence of inter‐pool movement. Rockpool depth, however, proved to be important in the blennies' movements. Our findings are among the first to prove that a mark‐recapture method can be successfully used to track intertidal fish movements. In particular, we showed that G. paganellus presents site fidelity in intertidal rockpools during its early ontogeny for a period of two to three months.     

Larval development of three blenniid species Aidablennius sphynx, Coryphoblennius galerita and Lipophrys canevai (Pisces: Blenniidae: Blenniini) in the western Mediterranean

The early stages of development of three blenniid species, Aidablennius sphynx (6.7–15.8 mm BL), Coryphoblennius galerita (4.3–13.9mm BL), and Lipophrys canevai (3.5–10.4 mm BL) are described from specimens collected in the western Mediterranean. The characteristics used for identification included meristic, developmental, morphological and pigmentation characters. Distinguishing characters of these species useful in differentiating them from other species of blenniids for which early stages are known in the Mediterranean are presented. Information on the occurrence of larvae and juveniles of these species off the Catalan coast (north-western Mediterranean) is also given.     

A method for long-term measurement of respiration in intertidal fishes during simulated intertidal conditions

Aquatic and aerial respiration of the amphibious fishes Lipophrys pholis and Periophthalmus barbarus were examined using a newly designed flow-through respirometer system. The system allowed long-term measurements of oxygen consumption and carbon dioxide release during periods of aquatic and aerial respiration. The M o ₂ of L. pholis , measured at 15° C, was 2·1 μmol O₂ g^–1 h^–1 during aquatic and 1·99 μmol O₂ g^–1 h^–1 during aerial exposure. The corresponding values of the M co₂ were 1.67 and 1.59 μmol O₂ g^–1 h^–1 respectively, giving an aquatic respiratory exchange ratio (RER) of 0·80 and an aerial RER of 0·79. The M o₂ of P. barbarus , measured at 28°C, was 4·05 μmol O₂ g^–1 h^–1 during aquatic and 3·44 μmol O₂ g^–1 h^–1 during aerial exposure. The corresponding values of the Mco₂ were 3·29 μmol CO₂ g^–1 h^–1 and 2·63 μmol CO₂ g^–1 h^–1 respectively, giving an aquatic RER of 0·81 and an aerial RER of 0·77. While exposed to air for at least 10 h, both species showed no decrease in metabolic rate or carbon dioxide release. The RER of these fishes equalled their respiratory quotient. After re-immersion an increased oxygen consumption, due to the payment of an oxygen debt, could not be detected.     

An assessment of the genotoxic impact of the Sea Empress oil spill by the measurement of DNA adduct levels in selected invertebrate and vertebrate species

The grounding of the Sea Empress oil tanker resulted in the release of 72,000 tonnes of crude oil into Milford Haven, Wales, UK. Our initial studies indicated that this contamination resulted in elevated levels of DNA adducts in one of the area's native marine species Lipophrys pholis [B.P. Lyons, J.S. Harvey, J.M. Parry, An initial assessment of the genotoxic impact of the Sea Empress oil spill by the measurement of DNA adduct levels in the intertidal teleost Lipophrys pholis, Mutat. Res. 390 (1997) 263-268]. These original studies were extended and the genotoxic impact of the oil contamination was investigated in the invertebrates Halichondria panicea and Mytilus edulis, along with the vertebrate fish species L. pholis, Pleuronectes platessa and Limanda limanda. DNA adduct levels were assessed in these species over a period of 2-17 months after the incident. The studies indicate differences in the impact of acute oil contamination upon vertebrate and invertebrate species. The oil contamination did not induce any detectable elevations in adduct levels in the invertebrate species H. panicea and M. edulis. In contrast, the oil contamination did appear to induce adducts in the vertebrate teleost species L. pholis, P. platessa and Lim. limanda. Despite some difficulties in sampling, the data obtained 12-17 months after the spill suggested that the affected species recovered from the oil contamination. While the studies indicate that the genetic impact of the oil contamination was less severe than might have been expected, it remains possible that the DNA adducts detected in the teleosts could lead to genetic changes in these species in the future.     

Comparative morphology of male reproductive systems in Mediterranean blennies (Blenniidae)

The male reproductive organs of 16 species of Mediterranean Blenniidae ( Aidablennius sphynx, Blennius ocellaris, Coryphoblennius galerita, Lipophrys adriaticus, L. canevae, L. dalmatinus, L. nigriceps, Parablennius gattorugine, P. incognitus, P. sanguinolentus, P. rouxi, P. tentacularis, P. zvonimiri, Paralipophrys trigloides, Salaria pavo and Scartella cristata ) consist of pairs of testes, testicular glands, spermatic ducts, and blind pouches. Three main types of accessory sex organs were found by comparing the external morphology of the male gonads. Differences between species were observed in the volume of the testicular gland in relation to the volume of the testis and in the size and length of the spermatic ducts, and blind pouches. The anatomy of the testicular glands of all species investigated do not differ. Each gland consists of ducts that appear to be tubules which terminate at the testis periphery on one side and at the spermatic duct on the other side. Contrary to previous claims, A. sphynx has no fat body in place of the testicular gland; the gland of this species was not distinguishable from that of the other species investigated. In the Lipophrys species, in P. trigloides , and in B. ocellaris , a transition zone between testis and testicular gland is present. The testicular blind pouches empty into the spermatic ducts, into the ureter, or separately on the genital papilla. In most species, the epithelium has no or low folds, while in S. pavo it possesses high folds that nearly fill the lumen of the blind pouches. The morphological results are discussed in connection with taxonomy, ecology, and behaviour of the fishes.     

Effects of temperature on the food processing of three species of seaweed-eating fishes from European coastal waters

Food consumption, gut transit times and assimilation efficiencies of the temperate fishes Lipophrys pholis and Parablennius sanguinolentus (Blenniidae) and Gobius cobitis (Gobiidae. when fed the green alga Ulva lactuca were determined at four temperatures in the laboratory. Food consumption increased and gut transit times decreased with increase in temperature in all three species. Assimilation efficiencies for algal carbon and nitrogen did not vary with temperature in G. cobitis . Carbon but not nitrogen assimilation efficiency varied with temperature in P. sanguinolentus , and both carbon and nitrogen assimilation efficiencies changed broadly with temperature in L. pholis , the most omnivorous of the three fishes. Assimilation efficiencies of all three species were generally within the previously reported ranges for herbivorous marine fishes. These species are thus clearly capable of extracting energy from algal food sources.     

Herbivore-induced infochemicals influence foraging behaviour in two intertidal predators

Herbivore-induced defences appear ubiquitous across most biomes and habitats. Yet the direct correlation between induced changes in host plant chemistry and the population dynamics of the herbivore remain untested in many systems. In plant–herbivore interactions in the terrestrial environment, indirect or tritrophic interactions appear a successful way in which changes in the host plant chemistry induced by prior herbivory can impact on herbivore populations via increased success of natural enemies. This set of interactions remains untested in the marine system. Here, we present work from experiments using orthogonal contrasts of plants with different prior treatments (control, mechanical damage or herbivory) and the presence or absence of herbivores on the foraging behaviour of a crab, Carcinus maenas, and a fish, Lipophrys pholis. These experiments were carried out using a novel flow-through flume, i.e. as a choice chamber supplied by turbulent water from independent cue sources. Our results show that in the Ascophyllum nodosum (plant)–Littorina obtusata (herbivore) system infochemicals from induced plants can directly influence predator foraging behaviour. L. pholis was attracted to the presence of a feeding L. obtusata, but was also more attracted to odours from herbivore-induced tissue than odours from mechanically damaged or naïve A. nodosum. C. maenas was more attracted to odours from herbivore-induced tissue compared to naïve tissue, regardless of the presence of L. obtusata. This is the first demonstration of such behavioural consequences of herbivore-induced changes in plants for marine systems.     

Habitat Utilization and Depth Distribution of Small Cryptobenthic Fishes (Blenniidae, Gobiesocidae, Gobiidae, Tripterygiidae) in Ibiza (Western Mediterranean Sea)

The paper describes the habitat utilization of small cryptobenthic fishes of the families Blenniidae (Lipophrys nigriceps, and Parablennius zvonimiri), Gobiesocidae (Apletodon incognitus, Gouania wildenowi, Lepadogaster candollei, and L. lepadogaster), Gobiidae (Chromogobius zebratus, Corcyrogobius liechtensteini, Didogobius splechtnai, Gammogobius steinitzi, Millerigobius macrocephalus, Thorogobius ephippiatus, T. macrolepis, and Zebrus zebrus), and Tripterygiidae (Tripterygion melanurus). Nine different habitats from the waterline to a depth of about 40m were examined. The depth distributions of the single species and co-occurrences with other cryptobenthic fish species are given.     

An initial assessment of the genotoxic impact of the Sea Empress oil spill by the measurement of DNA adduct levels in the intertidal teleost Lipophrys pholis

The Sea Empress oil spill resulted in the release of vast quantities of potentially genotoxic contaminants into the coastal environment of the county of Pembrokeshire (UK). We are at present attempting to determine the potential genotoxic impact of the incident upon the native marine species of the area. Here we describe the levels of DNA adducts in specimens of the intertidal teleost, Lipophrys pholis, exposed to extensive oil contamination as an indication of exposure to potential genotoxins. We detected elevated levels of adducts in L. pholis specimens from an area that underwent heavy oil contamination as compared to specimens from a clean reference area devoid of oil contamination. These preliminary studies indicated that the oil contamination induced DNA adducts in the L. pholis specimens, which could potentially cause genetic damage in this native marine species. Further studies are now required to assess the full extent of the genotoxic impact of the oil spill upon the Pembrokeshire area's native marine life.     

A comparative study of territoriality in intertidal and subtidal blennioids (Teleostei, Blennioidei)

This paper presents a comparative analysis of territoriality in three intertidal (Lipophrys pholis, Coryphoblennius galerita, Salaria pavo) and two subtidal (Tripterygion delaisi, Parablennius pilicornis) blennioid fishes. Focal-animal observations of males guarding eggs showed that: (i) intertidal species had smaller territories that were less frequently patrolled; (ii) in subtidal species feeding was limited to the territory, while in intertidal fishes a substantial proportion of the feeding acts occurred outside the defended area; (iii) intertidal species spent less time out of the nest and showed a lower level of locomotory activity; (iv) subtidal species were subjected to a higher number of territorial intrusions with more species intruding their territory, some of them potential egg predators; (v) subtidal species performed several water column displays that played a significant role in courtship, while in intertidal fishes these displays are almost absent. Signalling without the presence of a female was almost absent in intertidal species. It is argued that intertidal fishes minimize the time spent out of the nest and the loss of contact with the substrate, features that may be of high survival value in conditions of strong turbulence. Although intertidal species have a reduced time available for feeding, this may be compensated by lower levels of locomotory activity, territorial defense and risk of egg predation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Unusual testicular accessory organs, the testicular blind pouches of blennies (Teleostei, Blenniidae). Fine structure, (enzyme-) histochemistry and possible functions

The testicular blind pouches (TBPs) of Lipophrys canevae and Solaria pavo have an annual reproductive cycle with an active secretion phase during spawning, a regression phase correlated with necrotic processes during post-spawning, an inactive regeneration phase during inter-spawning and prespawning. As revealed by fine structure and enzyme-histochemistry the TBPs of L. canevae and S. pavo are involved in the production of steroid glucuronides, which possibly act as pheromones, Further functions are in the secretion of small amounts of sialomuciri and in lytic activity.     

Validation of otolith daily increments in early juveniles of shanny Lipophrys pholis

To assess the periodicity of micro‐increment formation in otoliths of Lipophrys pholis, 90 early juveniles were immersed in alizarin red S or tetracycline hydrochloride for 24 h and sacrificed after 10, 20 and 30 days. The number of micro‐increments viewed under light microscopy was significantly related to the duration of the experimental period, and the slopes of the linear regressions were not significantly different from 1. This study indicates that micro‐increments in sagittae were deposited daily and can be used as reliable sources of age information for L. pholis.     

Composition, structure and distribution of the ichthyoplankton in a Mediterranean coastal lagoon

The species composition and assemblage structure of the ichthyoplankton from the Mar Menor Lagoon in south‐east Spain are given. The fish larvae were sorted from zooplankton samples collected at 20 stations with a plankton net (50 cm mouth diameter and 500 μm mesh) during 36 surveys between February and December 1997. A total of 39 575 fish larvae representing 14 families, 22 genera and 36 species were identified. Gobiidae was the most dominant family (77·0%) followed by Blenniidae (19·4%) and Atherinidae (1·3%). The most abundant species were Gobius niger and Gobius paganellus , which accounted for 42·7 and 19·3% of the total respectively. These species were followed in order of relative abundance by Pomatoschistus marmoratus (13·9%), Parablennius pilicornis (9·4%), Lipophrys pavo (7·7%), Atherina boyeri (1·3%) and Parablennius tentacularis (1·3%). The high species diversity (2·0–2·8 bits individual⁻¹ for the annual diversity spectra at each sampling station) reflected a diverse assemblage of species. The main commercial species in the lagoon (Sparidae and Mugilidae) were poorly represented among the ichthyoplankton and they probably enter the lagoon on the bottom as recruits. Chlorophyll a concentration in the water column was the main factor explaining the seasonal variation in larval abundance. Spatial distribution of larvae was related to hydrographical circulation patterns in the lagoon and the movement of marine‐spawned larvae through the channels connecting the lagoon with the Mediterranean Sea.