The identification of Pomatoschistus minutus (Pallas) and Pomatoschistus lozanoi (de Buen) (Pisces, Gobiidae)

Sand gobies areextremely abundant in coastal waters and estuaries. Pomuroschistus lozunoi tends to be overlooked due to identification problems. This paper summarizes the characteristics to distinguish P. lozanoi from P. minutus and presents a new distinctive feature that allows identification of juveniles from 15-20 mm standard length onward. The known geographic distribution of P. lozanoi is extended northward to 53°30' N and southward to 40°N on the coast of Europe. In northern Europe P. loxmoi seems better adapted to the estuarine habitat than previously thought.     

Big is better: the importance of size for reproductive success in male Pomatoschistus minutus (Pallas) (Pisces, Gobiidae)

The sand goby, Pomatoschistus minutus , was used in aquarium experiments to study the importance of body size for the ability of males to gain access to nest sites and mates. When several male and female P. minutus were allowed to reproduce together, on average, half of the males built nests, and half of those males obtained eggs in their nests. Males with nests were significantly larger than males without nests, and nests with eggs belonged to males larger than the males with empty nests. In another experiment, when two males were competing for one nest, the largest male occupied the nest when both fish were put into the aquarium simultaneously. However, when the smaller male had been allowed to establish a nest before the larger male was introduced, the small male could usually retain the nest. Males with a body length < 50 mm did not build nests at all in the early part of the breeding season. In female choice experiments, no preference for larger males was found. Thus, male-male competition for nest sites and behavioural differences between different sized males seem to be the main factors influencing the non-random mating success in male 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.     

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 .     

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     

Development and characterization of nine polymorphic microsatellite markers in the sand goby Pomatoschistus minutus (Gobiidae)

A microsatellite‐enriched genomic library was constructed for the sand goby, Pomatoschistus minutus (Pallas 1770), and nine polymorphic DNA microsatellite markers of high quality were successfully optimized. Characterization of 96 individuals from the Vaccarès lagoon (France) showed moderate to high levels of polymorphism (two to 54 alleles). All the markers conformed to Hardy–Weinberg equilibrium and showed no evidence of null alleles, large allele dropout, stuttering and linkage disequilibrium between pairs of loci. These markers successfully amplify in three closely related species and can be employed to investigate population genetic structure and to clarify paternity in Pomatoschistus species.     

Effects of a low oxygen environment on parental effort and filial cannibalism in the male sand goby, Pomatoschistus minutus

In fish, brood cycling parental males sometimes eat some orall of their eggs, a behavior termed filial cannibalism. Wetested predictions of filial cannibalism models related to thecost of parental care in the male sand goby, Pomatoschistusminutus, by increasing the parental effort (fanning expenditure)through reduced levels of dissolved oxygen to 39% in an experimentalgroup, whereas a control group had fully saturated water. Malesshowed both full-clutch cannibalism and partial-clutch cannibalismin both treatments. Giving the males one to three females tospawn with, we found that small clutches were completely eatenmore often than were larger ones, whereas partial-clutch cannibalismwas not affected by clutch size. Although treatment did notaffect filial cannibalism, it did affect a male's energy statesuch that males in the low oxygen treatment lost more body fat,indicating a greater fanning effort. This shows that males inthe low oxygen treatment allocated more energy to the presentbrood, potentially at the expense of future reproductive success.Our study strongly suggests that filial cannibalism in malesand gobies represents a strategic life-history decision asan investment in future reproductive success, and is not triggeredby a proximate need for food necessary for the male's own survival.Furthermore, males in the low oxygen treatment built nests withlarger entrances, and were less likely to rebuild their nestsafter destruction. Presumably, this makes fanning easier butthe nest more vulnerable to predators, suggesting a trade-offbetween fanning and nest defense.     

Mate preference for multiple cues: interplay between male and nest size in the sand goby, Pomatoschistus minutus

Female mating preferences are often based on more than one cue.In empirical studies, however, different mate choice cues aretypically treated separately ignoring their possible interactions.In the current work, we studied how male body size and sizeof the male's nest jointly affect mate preferences of femalesand gobies, Pomatoschistus minutus. The females were givena binary choice between males that differed either in body sizeor size of their nest or both. We found that neither body sizenor size of the nest alone affected male attractiveness, buttogether these 2 cues had a significant effect. Specifically,large males were more popular among females when they had alarge nest than when they occupied a small nest. The resultssuggest that if interaction effects between multiple mate choicecues are not considered, there is a danger of ignoring or underestimatingthe importance of these cues in sexual selection by female choice.     

Effects of current and historic habitat fragmentation on the genetic structure of the sand goby Pomatoschistus minutus (Osteichthys,Gobiidae)

Habitat fragmentation is a major force that will influence the evolution of a species and its distribution range. Pomatoschistus minutus, the sand goby, has a North Atlantic–Mediterranean distribution and shows various level of habitat fragmentation along its geographic repartition. The use of mitochondrial sequences of the cytochrome b (cyt b) gene and two co‐dominant sets of nuclear markers (introns and microsatellites) allowed us to describe the relationships between P. minutus populations belonging to several different geographical regions of Europe and to assess the structure of populations inhabiting the Golfe du Lion, along the French Mediterranean coast. The present study confirms that the taxon located in the Adriatic Sea (Venice) should be considered as a distinct species, separated approximately 1.75 Mya. The comparison of P. minutus between the Atlantic and western Mediterranean coasts using polymorphic co‐dominant markers revealed that they belong to two demographically independent units, and thus could be considered as well as distinct species, more recently separated (0.3 Mya). The Pleistocene glaciations seem therefore to have played an important role in the diversification of this complex. Finally, at a regional scale in the Golfe du Lion, P. minutus appears to form a single huge homogeneous population. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 175–198.     

A new species of Pomatoschistus (Teleostei,Gobiidae): the Mediterranean's smallest marine fish

The new sand goby species Pomatoschistus nanus (Teleostei: Gobiidae) is described from the northern coast of the Levantine Sea (eastern Mediterranean Sea) based on both morphological and DNA barcoding data. The new species is the smallest fish in the Mediterranean Sea and may be distinguished from congeners by the following features: predorsal area, first dorsal‐fin base and breast naked; δ‐pore missing; anterior point of the suborbital row b not reaching level of posterior point of suborbital row d; slightly emarginated caudal fin and nape coloration pattern. DNA barcode data clearly discriminate Pomatoschistus spp. in the neighbour‐joining tree with an average of 17·7% interspecific K2P distance. The most closely related taxon to P. nanus sp. nov. is Pomatoschistus bathi and the most distantly related is Pomatoschistus tortonesei with 11·9 and 21·9% K2P distances respectively. Morphometric and genetic data are also provided for Pomatoschistus bathi.     

An electrophoretic study of the systematic relationships of some closely related goby species (Pisces, Gobiidae)

An electrophoretic study of genetic variation at 31 loci in each of nine closely related gobiid species was carried out in order to assess their systematic relationships. The species used were: Pomatoschistus canestrinii, P. lozanoi, P. marmoratus, P. microps, P. minutus, P. norvegkus, P. pictus, Knipowitschia caucasica, Neogobius melanostomus. Genetic distances ( D ) and identities ( I ) were calculated by Nei's method for all 36 pairwise comparisons and dendrograms were constructed.
The results show that the very similar and cross-fertile species pair, P. lozanoi and P. minutus , have the highest I value; P. norvegkus has high I values with both species. Pomatoschistus marmoratus and P. microps also display high genetic similarity with each other. The remaining Pomatoschistus species and Knipowitschia caucasica are broadly and moderately equidistant from each other and from the five species refferred to above, while Neogobius melanostomus is considerably more distant from all other species.
These findings essentially agree with the pattern of relationships established using classical taxonomic approaches. However, the observation that Knipowitschia caucasica is as similar to Pomatoschistus spp. as the latter are to each other suggests that the generic distinction of Knipowitschia could usefully be reconsidered.     

Temperature selection and avoidance in the sand goby,Pomatoschistus minutus (Pallas), collected at different seasons

Synopsis The sand goby from the Oslofjord, Norway, is extremely eurythermal. In spring and autumn it avoids temperatures lower than about 4° C, in summer 6° C. Acclimation did not influence the lower avoidance temperature, but the critical thermal maximum, the upper avoidance temperature and the temperature where the whole fish darkened (the darkening temperature) varied with acclimation and season. The darkening temperature is suggested to be the upper temperature limit with the greatest ecological significance. The fish, collected at different seasons darkened at between 19.5 and 22° C. In the field the fish is not found at 20° C and higher.The preference temperature varied with season and with acclimation temperature, generally with low precision. In May, the preference temperature was 13.5° C, that is higher than the ambient temperature of 10° C. In summer, the temperature in sampling locality and preference temperature was the same, 17 and 16.5° C respectively. In October, temperature preference was 7.5° C as compared to 9° C in the field. The variation is explained as a behavioural thermoregulation to direct the fish towards optimal conditions at any time.The seasonal variation in preference temperature can not be ascribed only to seasonal variation in temperature, that is an acclimation phenomenon, but other factors are operative as well, factors which will modify the temperature tolerance in the fish.     

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.     

To see in different seas: spatial variation in the rhodopsin gene of the sand goby (Pomatoschistus minutus)

Aquatic organisms living in a range of photic environments require specific mechanisms to tune their visual pigments. Maximum absorbance (λ_max) of retinal rods in populations of the marine demersal sand goby, ( Pomatoschistus minutus ; Gobiidae, Teleostei) correlates with the local optic environment. It has been shown that this is not regulated through a physiological response by exchanging the rhodopsin chromophore. To test for evolutionary adaptation, the sequence of the rhodopsin ( RH1 ) gene was analysed in 165 Pomatoschistus minutus individuals from seven populations across its distribution range. Analysis showed a high level of intraspecific polymorphism at the RH1 gene, including nonsynonymous mutations on amino acids, known as spectral tuning sites. Population differentiation at these sites was in agreement with the observed differentiation in λ_max values. Analyses of d _N/ d _S substitution rate ratios and likelihood ratio tests under site-specific models detected a significant signal of positive Darwinian selection on the RH1 gene. A strong discrepancy in differentiation was noticed between RH1 gene variation and the presumably neutral microsatellites and mitochondrial data. Samples did not cluster according to geographical or historical proximity with regards to RH1 , but according to the general photic conditions of the habitat environment of the sand goby. This study highlights the usefulness of sensory genes, like rhodopsin, for studying the characteristics of local adaptation in marine nonmodel organisms.     

Ionic basis for color changes in the iridescent cornea of the sand goby (Pomatoschistus minutus).

The iridescence from the cornea of the sand goby (Pomatoschistus minutus) occurs because of thin layer interference from the platelet-like cells in the stroma. It is suggested that ionic pumps across the epithelium control the water content in the stroma and thus the spectral reflection. A saline was perfused over goby eyes and simple ion manipulation was carried out to observe any changes in the iridescent characteristics. It was found that removal of Cl- and K+ ions reduced the peak reflected wavelength to the blue end of the spectrum, whereas Na+ had little effect. The removal of K+ also caused a dramatic change to the normal shift in reflected spectral intensity. The iridescence was also found to be sensitive to pH, and the buffer HEPES was detrimental to the cornea compared to controls. These results suggest similarities to amphibian and mammalian corneal hydration control.     

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.     

Rod opsin cDNA sequence from the sand goby (Pomatoschistus minutus) compared with those of other vertebrates.

The absorbance spectra of rods from the sand goby were measured by using microspectrophotometry. Analysis of the averaged spectra shows that the rod visual pigment has a maximum absorbance (lambda max) at approximately 501 nm. A sand goby retinal cDNA library was constructed and then screened with a partial sand goby rod opsin clone obtained by the polymerase chain reaction (PCR). The screening of the library yielded a full length rod opsin clone. The cDNA sequence and deduced amino acid sequence of this clone are compared with those of other vertebrate rod opsins.