Studies on scale regeneration in the Common goby, Pomatoschistus microps (Pisces)

The longevity of original scales and the process and consequences of scale regeneration in the euryhaline Common goby, Pomatoschistus microps (Kroyer), has been studied by monthly sampling of two populations. Scale regeneration comprises four stages: formation of a new growth centre, a regenerated area in the form of a wide central irregular area lacking concentric circuli: formation of short and widely spaced circuli at the outermost region of the regenerated area: a check in regenerated scale growth on reaching the size of the original scales: and the commencement of normal growth similar to that of the surrounding original scales. Growth of regenerated scales within 30 days was not sufficient to attain the same size of the original ones. The average percentage of regenerated scales in successive months during the life span of P. microps is about 20% in immature fish and sharply increases up to about 60% of the total scales during the breeding season. The correlation between sharp increases in regenerated scales and the breeding season is also confirmed on examining scales from two other gobioid species, Gobius paganellus and Gobius cobitis. It is suggested that the loss of a large number of scales may result in the check in body growth during the breeding season.     

Studies on scale structure in the common goby Pomatoschistus microps Krøyer

A histological study on scale structure in the common goby, Pomatoschistus microps (Krøyer) has established a similarity to other teleost scales, but yielded two new findings. First, the osseous layer of the scale was formed of successive and parallel calcified bands that may reflect a rhythm in scale growth. Second, extracellular matrix vesicles (100–200 nm) were identified by electron microscopy, at the calcifying area in the anterior growing edge of the scale; distribution of these vesicles corresponded closely to the pattern of scale calcification. A proposal on the mechanism of growth and calcification of scale has been made. The surface topography of the scale has also been examined by scanning electron microscopy. The raised edge of the circulus is formed of minute knobs, which may be used as criterion in fish taxonomy.     

The seasonal partitioning of energy in an estuarine fish, the common goby, Pomatoschistus microps Krøyer

The common goby, Pomatoschistus microps , is an abundant brackish water species which has an appreciable role, both as predator and prey, in the ecosystems of British estuaries. The partitioning of energy within each sex is described and discussed, and energy values are provided for the gonad, liver and carcass throughout almost the entire life of one year class. Partitioning of resources within the body at different times of the year shows considerable variation, not only with time but also between the sexes. In particular, the energy content of the male gonad during the spawning season is never more than 3.7% of that of the female. Important aspects of the biology of the species which relate to energy partitioning are maturation at an early age, high reproductive investment, high stochastic mortality, and death usually after a single breeding season.     

Alkaline phosphatase activity in the skin of the Common goby, Pomatoschistus microps (Krøyer), in relation to cycles in scale and body growth

A histochemical study of the distribution of alkaline phosphatase in the skin of the Common goby, Pomatoschistus microps , showed that the enzyme is present in the dermis, not in the epithelium, and is more pronounced around the growing edge of the scale and below the fibrillary plate. The intensity of the enzyme reaction is higher in regenerated scales. In conjunction with the phenology of P. microps , a method was adopted to measure the enzyme activity in the skin, including scales, for successive months over a period of two years. Enzyme activity displayed seasonal fluctuations which were closely related to both body and scale growth. Enzyme activity also reflected other phenological cycles during the life-span of P. microps .     

Phylogeography of the common goby, Pomatoschistus microps, with particular emphasis on the colonization of the Mediterranean and the North Sea

The phylogeographical patterns of a small marine fish, the common goby, Pomatoschistus microps, were assessed at 12 sites along the northeastern Atlantic coasts and the western Mediterranean Sea. A combination of two genetic markers was employed: cellulose acetate allozyme electrophoresis (CAGE) and sequence analysis of a 289 bp fragment of the mitochondrial locus cytochrome b. Both markers were congruent in revealing significant differences between samples (global FST = 0.247 for the allozymes and PhiST = 0.437 for the mitochondrial DNA data) and a pattern of isolation-by-distance. Phylogeographical analyses yielded a shallow branching structure with four groups. Three of those were confined to the Atlantic basin and showed a star-like pattern. The fourth group contained a central haplotype occurring at the edges of the species' distribution, accompanied by a few more rare variants, which were restricted to the Mediterranean Sea. A genetic break was observed around the British Isles, with distinct haplotypes dominating at either side of the English Channel. A significantly negative correlation between the degree of genetic diversity and latitude was recorded both for mitochondrial DNA (mtDNA) and allozymes in the Atlantic basin. Gene flow analysis suggested that recolonization of the North Sea and the coasts of western Scotland and Ireland may have taken place from a glacial refugium in the Southern Bight of the North Sea. These results are discussed in the perspective of possible postglacial migration routes of marine fish along the northeastern Atlantic coasts.     

Reproductive effort and efficiency in the female common goby, Pomatoschistus microps (Krøyer) (Teleostei: Gobioidei)

Reproductive effort and efficiency are terms which together help to describe the reproductive investment of spawning fish. In this paper they are applied to a small and abundant inshore fish, the common goby, Pomatoschistus microps (Kroyer), based on data collected from the wild, and by modelling likely breeding patterns. Wet calorimetry was used to determine the energy content offish tissue, and this allowed reproductive effort to be expressed as both an energy and a weight ratio of gonad to somatic body. The gonadosomatic index (GSI) calculated as an energy ratio was greater than that calculated as a dry weight ratio. For ripe females the GSI (dry weight) at the onset of breeding was 67.5%. Theoretical and observed values of the maximum number of egg batches laid by a single female during a 16-week breeding season are 9-10, and this corresponds to a total caloric investment of almost 12 kJ. Estimates of the reproductive efficiency, using the dietary energy content during the breeding season, range from 28.2% to a maximum of 44.0%. These values are among the highest calculated for a fish species, and are in keeping with the long and intensive breeding strategy observed in the wild population.     

Parasitological diversity of the common goby, Pomatoschistus microps (Krøyer, 1838), in estuarine systems along the Portuguese coast

Parasitological diversity of the common goby Pomatoschistus microps along the Portuguese coast was studied based on samples collected using beam trawls in five coastal estuaries (Ria de Aveiro, Tagus, Sado, Mira and Guadiana estuaries) between May and July 2006. The classes Digenea, Cestoda, Nematoda and phylum Mastigophora were found, with Digenea Cryptocotyle sp. and Lecithochirium musculus being the most abundant taxa. Prevalence and mean abundance of Cryptocotyle sp. were highest in Ria de Aveiro (74%) and Mira estuary (28.05), respectively, whereas L. musculus values of both indices were highest in Sado estuary (prevalence = 64%; mean abundance = 11.77). Values varied along the Portuguese coast; the discriminant analysis based on the number of parasites showed some overlap between estuaries, although successfully separating the Sado and Mira estuaries. Overall, variations in estuaries were explained by different environmental pressures, leading to differences on host abundance and life-history of the parasites. Feeding behaviour and habitat selection therefore play a crucial role in shaping the biology, ecology and parasite assemblages of the common goby along the Portuguese coast.     

Female preference for preferred males is reversed under low oxygen conditions in the common goby (Pomatoschistus microps)

Female preference for males that already have eggs in theirnest has beenreported in many fish species. The presence ofeggs may provide a cue forcopying the choice of previous femalesand may indicate that high-quality carewill be available. Consistentwith a review of 13 studies, we found thatfemale common gobies(Pomatoschistus microps) preferred to spawn withmales thathad already been chosen by females and whose nests thereforealreadycontained eggs. However, this preference was reversed underconditionsof low dissolved oxygen. We would not expect thisreversal if the secondfemale were using eggs as a signal ofmale genetic attractiveness to otherfemales unless the benefitswere outweighed by direct selection. The reversalalso couldnot be explained by differences in active courtship by males,ascourtship rates did not differ under low oxygen between maleswith or withouteggs. Low oxygen conditions corresponded witha nearly threefold increase inmale ventilation of eggs anda reduction in time spent near a selectingfemale. The reversalis therefore most likely due to females avoiding malesthatwould be unable to meet the demands of care of a second clutchunder lowoxygen conditions. Thus, an abiotic feature of theenvironment revealsplasticity of female choice, consistentwith hypothesized changes in benefitsof mating with preferredmales.     

Phosphoglucose isomerase polymorphism in the common goby, Pomatoschistus microps (Krøyer) (TeleosteirGobiidae), around the British Isles

Phosphoglucose isomerase (PGI) polymorphism has been investigated by horizontal starch-gel electrophoresis in the common goby, Pomatoschistus microps (Krøyer), from 21 localities around the British Isles and at Roscoff, France. Sixteen PGI phenotypes were observed, consistent with occurrence of four alleles at each of the two loci (A,B). In the Teign estuary at Shaldon, Devon, seasonal fluctuation in PGI-A allele frequencies was accompanied in some months by departure from Hardy-Weinberg equilibrium. At this locality, a significant change in PGI-A allele frequencies was found with increasing modal age of sample (within a year class) but not between year-classes over eight years. Similar stability is noted for other areas. Major alleles (PGI-A2, A3; PGI-B2, B3) were found at all localities, but with marked differences in proportions. Atlantic coast populations tended to have higher A2 and B2 frequencies than those of the North Sea and English Channel, with Irish Sea populations having high A2 but low B2. Historical and selective causes for this geographical variation are discussed. A significant correlation exists between the frequencies of the major PGI-A alleles and maximum monthly mean sea temperature for the same area.     

Structure,growth and production of a remarkably abundant population of the common goby,Pomatoschistus microps (Actinopterygii: Gobiidae)

The common goby, Pomatoschistus microps, is a relevant species from estuarine food webs, playing important roles as predator of polychaetes and crustaceans and as prey for larger fishes and crustaceans. The Minho estuary (NW Portugal) is a relatively well-preserved and productive system. To assess the population structure and production of P. microps in this estuary, monthly samples were undertaken in three different areas along an estuarine gradient in the lower estuary. The density of P. microps varied considerably among seasons and sampling stations, with higher densities occurring in summer and autumn. The lowest densities were found closer to the sea. In general, the density of females was higher than the density of males in all sampling stations, while juveniles were more abundant within a salt marsh area. Compared with other European estuaries, our data showed a remarkable higher density and production values of P. microps. This may be related to the high freshwater input and the low salinities found in this estuary. In addition, we hypothesize that the lower density of the sympatric species P. minutus and the high availability of bivalve shells observed in the Minho estuary may have also contributed to the present results, once P. minutus and P. microps often display a diet overlap and the bivalve shells are crucial for the common goby reproduction.     

Asymmetry, reproductive success and parasitism of Pomatoschistus microps in a French lagoon

There was an allometry between length of nesting male Pomatoschistus microps and the size of the nest. There was also a positive relationship between parasite load and pectoral fin asymmetry. There was no marked relationship between the number of eggs in the nest and parasitism or level of asymmetry. These results are discussed in terms of energy allocation and fish reproductive success.     

Long-term monitoring on the occurrence of a myxosporean parasite Kudoa camarguensis (Myxosporean) on the common goby (Teleostei, pisces) Pomatoschistus microps.

The evolution of a host-parasite system composed of Pomatoschistus microps-Kudoa camarguensis was investigated in the Vaccarès lagoon (Rh?ne river Delta, France) from 1993 to 1997. During this long-term monitoring, centennial flooding of the Rh?ne river occurred, leading to an inrush of about 110 million m3 of freshwater in the Vaccarès lagoon. The salinity drastically dropped from 14 to 5 g l(-1) in 1 wk. We observed that the annual prevalence and abundance of the myxosporean parasite decreased from 12.18 in 1993 to 3.7% in 1997 and from 1.10 in 1993 to 0.27 in 1997, respectively. Here, we discuss the possible reasons for the rapid decline of this host-parasite system following the flood.     

Behaviour and success of sneaker males in the sand goby, Pomatoschistus minutus

In this study we investigated in laboratory conditions the presence of alternative reproductive tactics in the sand goby and describe proximate factors affecting their expression and success. We describe the reciprocal interactions of resident males, females, and sneaking males. The pre-spawning phase proved to be important for successful nest intrusions by sneakers. The number of sneakers had no effect on the frequency of successful intrusions. When small males had exclusive access to nest sites, they built a nest and courted females, showing a full behavioural repertoire. The intensity of courtship was, however, strongly positively correlated with body size. Using microsatellite DNA markers we assessed paternity shares of territorial and sneaker males in a subset of all replicates. Following successful nest intrusion sneaker males fertilised 5–10% of the eggs. Our interpretation of the results is that sneaking in the sand goby is a conditional tactic, one that is less successful than the normal nest guarding behaviour, at least for one spawning event. Received in revised form: 9 March 2001 Electronic Publication     

First record of Kudoa (Myxosporea) in Pomatoschistus microps in the Rhône Delta, France

Myxosporidiosis due to Kudoa sp. has been recorded in gobies Pomatoschistus microps , collected from the Etang du Vaccarès (Rhône Delta, France).     

Reproductive biology of a Red Sea goby, Silhouettea aegyptia, and a Mediterranean goby, Pomatoschistus marmoratus, in Lake Timsah, Suez Canal

The reproductive biology of the Red Sea goby Silhouettea aegyptia , and Mediterranean Pomatoschistus marmoratus , has been studied in Lake Timsah, Suez Canal. Male: female sex ratio was 1: 1 7 in S. aegyptia and 1: 2 7 in P. marmoratus . Size at first maturity was 20 mm ( s.l. ) in S. aegyptia and 24 mm in P. marmoratus . The breeding season of S. aegyptia extended from March to September with most gonadal activity in May, whereas in P. marmoratus , it stretched from. November until the following April with a peak in February. Hepatosomatic index for both species exhibited monthly fluctuations and an inverse correlation with gonadosomatic index was evident. Oocyte size-frequency distribution indicated several modes at 400, 600 and 800 μm during the breeding season. Fecundity of S. aegyptia ranged from 152 to 782 with an average of 468 eggs, whereas in P. marmoratus it varied between 293 and 1300 with an average of 603 eggs. Fecundity was related curvilinearly to fish length, body weight and ovary weight. Recruitment of young fish to the adult populations occurred in late July and early August for S. aegyptia and during March for P. marmoratus . Life history pattern indicated that both species of gobies are successful colonists of Lake Timsah, Suez Canal.     

Size-assortative nest choice in the absence of competition in males of the sand goby,Pomatoschistus minutus

Synopsis Size-assortative choice of nest site is commonly found among fish with paternal care. This pattern can be explained either by the superior competitive ability of large males for preferred large nests, or by a choice of nest size that is related to the male's own body size. Choice of nest size was studied in males of a small marine fish, the sand goby Pomatoschistus minutus, by providing artificial nest sites in laboratory and field experiments. A positive correlation between nest size and male size, indicating a size-assortative nest choice process, was found in both aquarium and field experiments. This is argued to be the outcome of a preference for an optimal nest size and not to be due to competition, as (a) a surplus of nest sites was available to the individuals in nature; (b) in the aquarium experiments, single males of all sizes, when offered a choice between a smaller and a larger nest site, showed a preference for nests in proportion to their body size and not for the largest available nests; and (c) in the aquaria, when single males were offered only a small or only a large nest site, small males abstained from building nests in the large sites, whereas in the small nest sites males of all sizes built nests. My conclusion is that size assortative nest choice occurred in the absence of competition, which thus is not a necessary condition for size-assortative nesting, contrary to what is commonly assumed in many studies. This preference may represent a compromise (or trade-off) between the advantage of a nest large enough to accommodate many eggs and a smaller one with lower maintenance and defence costs.     

Genetic architecture of the marbled goby Pomatoschistus marmoratus (Perciformes, Gobiidae) in the Mediterranean Sea

The marbled goby Pomatoschistus marmoratus, a species inhabiting coastal Mediterranean lagoons, has been studied by measuring its mitochondrial DNA variation. This analysis revealed a Mediterranean west vs east split and, subsequently, an eastern differentiation among the Libyan-Tunisian Gulf, the Adriatic Sea and the Aegean Sea. The high cohesion between the samples collected in the vast area of western Mediterranean contrasts with the genetic mosaic of the more sub-structured eastern Mediterranean. This western homogeneity can not yet be fully explained even if a human-mediated migratory flow, due to a maritime traffic, has been posited. The pattern in the eastern basin revealed a genetic architecture possibly due to the non-migratory habit of the gobid. Within this perspective, the role of the Mediterranean lagoon habitat should be related to how much it amplifies the effects of historical (e.g. past sea-level changes) and environmental (e.g. present-day hydrographic regime) processes as regards the genetic structure of the inhabiting species.     

Egg presence, egg loss, and female mate preferences in the sand goby (Pomatoschistus minutus)

We studied the effect of egg presence on female mate choicein a fish with paternal care. Females who were allowed a freechoice between two males mated within a shorter time than femaleswho were randomly assigned to a particular male. When a secondfemale was allowed to choose among the males, she preferredthe same male as the previous female. This result shows thatfemales are concordant in their mate choice. When the initialfemale was randomly assigned to mate with one of two males (forcedchoice), the second female mated randomly with respect to thefirst one. Thus females do not prefer males with eggs. If theinitial female was given a free choice, but the eggs were removedfrom the chosen male, the test female mated randomly. When boththe males initially had mated but one randomly determined male'seggs were removed, the test female preferred the male who wasstill guarding eggs. These experiments show that females avoidspawning in unsuccessful nests. When the females in the freechoice/egg removal experiment mated with the unsuccessful malethere was a considerably bigger size difference in favor ofthis male than when the females mated with the other male. Weconclude that female sand gobies show clear mate preferences,but that they do not prefer males with eggs over males withouteggs. They do, however, avoid mating with males guarding unsuccessfulnests. We therefore suggest that egg loss could be an importantfactor selecting for egg preference.     

Age determination in Pomatoschistus mi nut us (Pallas) and Pomatoschistus microps (Kreyer) (Pisces: Gobiidae) from the upper Tagus estuary, Portugal

The use of scale inspection as a means of age-determination in Pomatoschistus microps and Pomatoschistus minutus in the Tagus estuary is confirmed, but annulus formation pattern is different from that described for P. microps in the area of the British Isles (Miller, 1975). In Britain, two annuli are laid down on the scales each year, but in the Tagus, only one annulus is formed. One possible reason for this difference is the fact that in the Tagus the peak breeding season and the lowest water temperatures coincide. Therefore, the two causes of annulus formation coexist at the same time of year.
Population structure in both species is dominated by the 0-group, which appears earlier (April/May) than reported for other areas. Maximum age seems not to exceed 26 months in P. microps and 32 months in P. minutus .     

Tolerance, behaviour and oxygen consumption in the sand goby, Pomatoschistus minutus (Pallas), exposed to hypoxia

Behavioural and metabolic reactions of Pomatoschistus minutus (Pallas) exposed to various degrees of hypoxia were studied. At 15°C and 20‰ mortality was 50% at 15.2% oxygen saturation. Avoidance and oxygen saturation showed a linear inverse relationship. At levels lower than 60% saturation increased activity occurred; avoidance was significant at 30% saturation. Active, routine and standard MO₂ correlated linearly with weight at 6 and 15°C (salinity = 19‰). During hypoxia at 15°C routine MO₂ rose significantly at 60–50% and 40–30% saturation expressed either as MO₂ during longer periods at night or MO₂ at shorter intervals during the day. Standard MO₂ was unaffected by hypoxia at 15°C. Haemoglobin concentration was significantly increased when P. minutus was acclimatized to 35% saturation.