“A cleaner break”: Genetic divergence between geographic groups and sympatric phenotypes revealed in ballan wrasse (Labrus bergylta)

Capture and long‐distance translocation of cleaner fish to control lice infestations on marine salmonid farms has the potential to influence wild populations via overexploitation in source regions, and introgression in recipient regions. Knowledge of population genetic structure is therefore required. We studied the genetic structure of ballan wrasse, a phenotypically diverse and extensively used cleaner fish, from 18 locations in Norway and Sweden, and from Galicia, Spain, using 82 SNP markers. We detected two very distinct genetic groups in Scandinavia, northwestern and southeastern. These groups were split by a stretch of sandy beaches in southwest Norway, representing a habitat discontinuity for this rocky shore associated benthic egg‐laying species. Wrasse from Galicia were highly differentiated from all Scandinavian locations, but more similar to northwestern than southeastern locations. Distinct genetic differences were observed between sympatric spotty and plain phenotypes in Galicia, but not in Scandinavia. The mechanisms underlying the geographic patterns between phenotypes are discussed, but not identified. We conclude that extensive aquaculture‐mediated translocation of ballan wrasse from Sweden and southern Norway to western and middle Norway has the potential to mix genetically distinct populations. These results question the sustainability of the current cleaner fish practice.     

Age, growth and feeding in the ballan wrasse Labrus bergylta Ascanius 1767

A study of the age, growth and feeding of the protogynous wrasse Labrus bergylta (Teleostei: Labridae) was made on 25 male and 316 female specimens taken from the southern coasts of the Isle of Man from October 1972 to October 1974. Age determination and back-calculations for length were made from the opercular bones. The oldest male and female specimens were 29 and 25 years old respectively. The growth curves showed only a slight increase in growth rate after sexual inversion, and a pattern of slow irregular increase, with no pronounced levelling off in old fish. The time of sexual inversion varied widely: intersexual specimens of 5 and 14 years were found. The youngest functional male was 6 years old. The weight/length relationships of males and females were very similar and the following equation, derived from all individuals, was used to calculate weights for age utilizing the back-calculated lengths:
The mouth, pharyngeal teeth and gut length to fish length ratio (0.622) suggested an omnivorous diet. Analysis of gut contents confirmed this, with decapod Crustacea, isopods and molluscs the dominant food items. There were seasonal changes in feeding intensity and composition of the diet, related to temperature changes, onshore-offshore migration patterns, availability of food items, and changes in habitat associated with maturation.     

Isolation,identification and characterisation of ballan wrasse Labrus bergylta plasma pigment

This study confirmed that observations of blue‐green colouration in plasma fractions of the ballan wrasse Labrus bergylta were caused by the linear tetra‐pyrrole biliverdin and that the molecule was of the physiologically relevant IXα isomer. Accumulation appears driven by chromogenic association with an unknown protein moiety which precludes enzymatic reduction and would suggest active management. It was demonstrated that the pigment did not fluctuate relative to ontogeny, or indeed binary gender in the species of interest, but mobilisation and depletion in the subset of individuals undergoing sex change at the time of study supports a potential association with gender inversion processes. It is of note that although biliverdin does have some effect on external colouration, the evidence is indicative that crypsis is a supplementary function thus other factors must be considered.     

Gender distribution,sexual size dimorphism and morphometric sexing in ballan wrasse Labrus bergylta

In wild ballan wrasse Labrus bergylta, mass–length relationships were not different between genders, and positive allometry was found in the mixed‐gender population. Male‐biased sexual size dimorphism was significant and the most effective morphometric method for sexing L. bergylta outside of the species spawning window used body mass (M_B in g), total body length (L_T in mm) and Fulton's condition factor (K) as discriminant variables to predict gender with 91% accuracy. The discriminant score (S_D) of a specimen can be calculated as S_D = 0·01 M_B ? 0·016 L_T ? 3·835 K + 6·252 to predict its gender as female or male if S_D is < 1·459 or S_D is > 1·504, respectively. There was a potential trend towards earlier sexual inversion compared to previous studies at comparable latitudes. Sex change is a phenotypically plastic trait under social control in haremic fishes and should be monitored in increasingly exploited L. bergylta.     

Interannual fine-scale site fidelity of male ballan wrasse Labrus bergylta revealed by photo-identification and tagging

The site fidelity of ballan wrasse Labrus bergylta was studied using photo-identification and external tagging. Five male individuals were observed to defend the same small territory composed of a few rocks during several reproductive seasons spanning 2 to 15 years. These results provide one of the strongest indications of long-term very fine-scale site fidelity in marine fishes.     

Natural sex change in the temperate protogynous Ballan wrasse Labrus bergylta

Wild Ballan wrasse Labrus bergylta were sampled monthly over 2 years in western Norway to identify the natural process of sex change in this species. Light microscopy of standard histological‐stained and immunohistochemistry‐treated gonad tissue showed that spermatogonial germ cells tended to proliferate around the periphery of the lamellae before filling into the slowly receding, apoptotic central areas of the lamellae. Sex change occurred following the breeding season. From July to September, fish were most often in an early state of gonadal transition (ET), characterized by degenerating previtellogenic oocytes and pockets of proliferating spermatogonia in the germinative epithelia. The majority of fish with late transitional gonads, that were typically dominated by spermatogenic cells, developing efferent ducts and the beginning of lobule formation, were found between October and November. Sex steroid profiles of fish representing the different sexual phases showed that breeding females had the highest concentrations of 17β oestradiol (E₂) and the lowest concentration of 11 ketotestosterone (11KT). Concentrations of E₂ decreased greatly in ET fish at the beginning of sex change and remained low in all subsequent phases. The opposite trend was demonstrated in 11KT profiles. Initial‐phase female fish had minimal concentrations of 11KT, but these increased during subsequent transitions. Sex change occurred most often in fish 34–41 cm total length (L_T) and the median of fish in the size‐frequency overlap of female and male fish was 36 cm L_T.     

The reproductive cycle of female Ballan wrasse Labrus bergylta in high latitude,temperate waters

This 2 year study examined the reproductive cycle of wild female Ballan wrasse Labrus bergylta in western Norway as a precursor to captive breeding trials. Light microscopy of ovarian histology was used to stage gonad maturity and enzyme‐linked immuno‐absorbent assay (ELISA) to measure plasma concentrations of the sex steroids testosterone (T) and 17β‐oestradiol (E₂). Ovarian recrudescence began in late autumn to early winter with the growth of previtellogenic oocytes and the formation of cortical alveoli. Vitellogenic oocytes developed from January to June and ovaries containing postovulatory follicles (POF) were present between May and June. These POF occurred simultaneously among other late maturity stage oocytes. Plasma steroid concentration and organo‐somatic indices increased over winter and spring. Maximal (mean ±s.e .) values of plasma T (0·95 ± 0·26 ng ml^?1), E₂ (1·75 ± 0·43 ng ml^?1) and gonado‐somatic index (I_G; 10·71 ± 0·81) occurred in April and May and decreased greatly in July when only postspawned fish with atretic ovaries occurred. Evidence indicates that L. bergylta are group‐synchronous multiple spawners with spawning occurring in spring and peaking in May. A short resting period may occur between late summer and autumn when previtellogenic oocytes predominate and steroid levels are minimal.     

Embryonic development in Erythrocebus patas

The developmental stages of 12 Erythrocebus patas embryos, ranging in gestational age from 30 to 50 days, is described. The pattern of embryogenesis in E. patas closely parallels the anatomic characteristics of human and other nonhuman primate embryos between stages 12 and 23. However, there is a delay in development in E. patas similar to that observed in human embryos which differs from the macaques and baboons. This temporal difference in the embryonic period is an important factor in the design and analysis of early pregnancy studies in this species.     

Uncertainty in territory quality affects the benefits of usurpation in a Mediterranean wrasse

Individuals should defend sites when the expected benefits ofthe territory exceed the cost of defense. However, if territoryqulaity is unpredictable or difficult to assess, the expectedpattern of territorial behavior is less clear. In a Mediterraneanwrasse, Symphodus ocellatus, mating success is skewed with 2%of nesting males getting more than 20% of the spawning success.Yet, variation in mating success is not explained by any knownphysical characteristic of males or their territories. Instead,females prefer nests with a recent history of mating successbecause males are less likely to desert the offspring she leavesbehind. Thus, territory quality is transient and determinedby interactions between the sexes. I measured the frequencyof territorial takeovers and the uncertainty in mating successamong days at a nest. Observations indicated that S. ocellatusmales usurped their neighbor's successful nests when males wereunsuccessful and larger than their successful neighbor. Sitesthat achieved mating success had a significantly higher probability(0.84) of remaining sucessful between consecutive days thanunsuccessful territories had of becoming successful (0.30).Unsuccessful males obtained higher and more certain fitnessreturns if they usurped a successful neighbor's territory. Interactionswithin and between the sexes drive uncertainty in success, whichinfluences territorial behavior in this species.     

双斑蟋的胚胎发育

为了从组织胚胎学角度探究昆虫胚胎发育过程,以双斑蟋Gryllus bimaculatus de Geer的卵为实验材料,通过观察、记录蟋蟀胚胎每一天的形态变化并使用显微摄影方法记录胚胎发育过程,对蟋蟀卵胚胎发育全过程进行系统的观察和研究。根据胚胎形态的发育特点,可将整个胚胎发育过程分为7个阶段:卵裂期、囊胚期、原肠胚、无节幼体期Ⅰ、无节幼体期Ⅱ、无节幼体期Ⅲ、无节幼体期Ⅳ。经历14天,蟋蟀的头部、触角、3对足、尾部、腹部及背部都发育完全,整个胚胎发育随之结束。     

Demographic history has shaped the strongly differentiated corkwing wrasse populations in Northern Europe

Understanding the biological processes involved in genetic differentiation and divergence between populations within species is a pivotal aim in evolutionary biology. One particular phenomenon that requires clarification is the maintenance of genetic barriers despite the high potential for gene flow in the marine environment. Such patterns have been attributed to limited dispersal or local adaptation, and to a lesser extent to the demographic history of the species. The corkwing wrasse (Symphodus melops) is an example of a marine fish species where regions of particular strong divergence are observed. One such genetic break occurred at a surprisingly small spatial scale (F_ST ~0.1), over a short coastline (<60 km) in the North Sea‐Skagerrak transition area in southwestern Norway. Here, we investigate the observed divergence and purported reproductive isolation using genome resequencing. Our results suggest that historical events during the post‐glacial recolonization route can explain the present population structure of the corkwing wrasse in the northeast Atlantic. While the divergence across the break is strong, we detected ongoing gene flow between populations over the break suggesting recent contact or negative selection against hybrids. Moreover, we found few outlier loci and no clear genomic regions potentially being under selection. We concluded that neutral processes and random genetic drift e.g., due to founder events during colonization have shaped the population structure in this species in Northern Europe. Our findings underline the need to take into account the demographic process in studies of divergence processes.     

Parasites of wrasse (Labridae) in inshore waters of west Scotland stocked as cleaner fish of sea lice (Caligidae) on farmed Atlantic salmon

The prevalence, abundance and intensity of protistan and metazoan parasites were examined in the labrid fishes Ctenolabrus rupestris, Centrolabrus exoletus, Crenilabrus melops, Labrus bergylta and L. mixtus in west Scotland. The 22 parasite species identified do not appear to be a major threat to farmed Atlantic salmon. Few mortalities of wrasse stocked on salmon farms were due to resident parasites or to transfers from salmon.     

Influence of batch-specific biochemical egg characteristics on embryogenesis and hatching success in farmed pikeperch

Low and variable egg quality remains a major issue in aquaculture impeding a reliable and continuous supply of larvae, particularly in emerging species, such as pikeperch, Sander lucioperca. We assessed the influence of batch-specific egg parameters (fatty acid (FA) profiles, cortisol content) on embryo life-stages until hatching (survival at 2, 24, 48, 72 h post fertilization (hpf), hatching rate) in an integrated study under commercial hatchery conditions (44 egg batches). Embryo mortality was elevated until 48 hpf (average 9.8% mortality between 2 and 48 hpf). Embryos surviving until 48 hpf were very likely (98.5%) to hatch successfully. The inherent egg FA composition was variable in-between batches. Total FA content ranged form 66.1 to 171.7 µg/mg (dry matter) total FA. Whereas specific FA,18 : 0 and 20 : 5(n-3) (eicosapentaenoic acid) of the polar fraction and the ratio of 22 : 6(n-3) (docosahexaenoic acid) to 20 : 5(n-3) within the neutral fraction, were significantly correlated with early embryo development, contents of the respective FA did not differ between high (>90% hatching rate), mid (70% to 90% hatching rate) and low (<70% hatching rate) quality egg batches. Late embryo development and hatching were relatively independent of the FA profiles highlighting stage-dependent influences especially during early embryogenesis. Cortisol levels ranged from 22.7 to 293.2 ng/ml and did not directly explain for mortalities. However, high cortisol was associated with a lower content of specific FA, in particular highly unsaturated FA. These results demonstrate the magnitude of inter-individual differences in the batch-specific biochemical egg composition under stable hatchery conditions and suggest a stress-mediated lack of essential FA, which in turn affects early embryo survival. Surprisingly, embryos are able to cope well with a broad range of inherent egg parameters, which limits their predictive potential for egg quality in general. Still, specific FA profiles of high quality egg batches have potential for formulating species-specific broodstock diets and improving reproductive management in pikeperch.     

Embryonic development of the endangered Caspian brown trout,Salmo trutta caspius (Kessler, 1877)

The Caspian brown trout, Salmo trutta caspius is an endangered species of economic value and better knowledge of its biology will contribute to conservation measures. In the present study we characterized its early development to identify critical embryonic stages, by observing embryos and larvae from a broodstock maintained in captivity incubated at 9 ± 1°C. Morphological analysis of landmark stages of embryonic development revealed that first cleavage occurred at 189 degree‐h, a many‐celled blastoderm formed at 62 degree‐days (dd), a germ ring at 72 dd, the neural keel stage at 88 dd, the blastopore closed at 137 dd, the eye became pigmented at 188 dd and the larvae hatched at 373 dd. The detailed timing of these landmark features will help development of strategies aiming at increased aquaculture production of this species.     

Functional morphology of embryonic development in the Port Jackson shark Heterodontus portusjacksoni (Meyer)

The oviparous Port Jackson shark Heterodontus portusjacksoni embryo has a long incubation of 10–11 months during which it undergoes major morphological changes. Initially the egg capsule is sealed from the external environment by mucous plugs in either end of the capsule. Four months into incubation, the egg capsule opens to the surrounding sea water. Fifteen stages of development are defined for this species, the first 10 occur within the sealed capsule, the remaining five after capsule opening to hatching. The functional significance of major definitive characters such as circulation within the yolk membrane and embryo, rhythmic lateral movement of the embryo, external gill filaments, heart activity, internal yolk supplies, egg jelly and the significance of the opening of the egg capsule are described. The egg jelly in the sealed capsule functions to mechanically protect the embryo during early development, however, it eventually creates a hypoxic environment to the embryo as the available oxygen is used up. This generates several physiological challenges to the developing embryo. It is able to overcome these problems by morphological changes such as increasing the effective surface area for gaseous exchange with the development of external gill filaments, fins and extensive circulation in both the embryo and attached external yolk sac. These adaptations become limiting as the embryo grows and respiratory needs outweigh the available oxygen. At this time, the mucous plugs dissolve and the capsule becomes open to the external environment.     

Prepared for the future: A strong signal of evolution toward the adult benthic niche during the pelagic stage in Labrid fishes

The morphology of organisms reflects a balance between their evolutionary history, functional demands, and biomechanical constraints imposed by the immediate environment. In many fish species, a marked shift in the selection regime is evident when pelagic larvae, which swim and feed in the open ocean, settle in their adult benthic habitat. This shift is particularly dramatic in coral‐reef fishes, where the adult habitat is immensely complex. However, whether the adult trophic ecotype affects the morphology of early‐life stages is unclear. We measured a suite of 26 functional‐morphological traits in the head and body of larvae from an ontogenetic series of 16 labrid species. Using phylogenetic comparative methods, we reconstructed the location of adaptive peaks of larvae whose adults are associated with different trophic ecotypes. We found that the morphospace occupation in these larvae is largely driven by divergent adaptations to the adult benthic habitats. The disparity between adaptive peaks is achieved early and does not monotonically increase with size. Our findings thus refute the notion that larvae rapidly acquire the trophic‐specific traits during a metamorphic period immediately prior to settlement. This early specialization might be due to the highly complex musculoskeletal system of the head that cannot be rapidly modified.     

The association between frequency of thiabendazole treatment and the development of resistance in field isolates of Ostertagia spp. of sheep

Martin P. J., Anderson N., Lwin T., Nelson G. and Morgan T. E. 1984. The association between frequency of thiabendazole treatment and the development of resistance in field isolates of Ostertagia spp. of sheep. International Journal for Parasitology14: 177–181. Isolates of Ostertagia spp. were obtained from grazing sheep 3,4 and 5 years after nil, planned (five per year) and regular (3-weekly) treatments with thiabendazole (TBZ). Levels of resistance to TBZ were measured by an in vitro egg hatch assay and a controlled anthelmintic efficiency assay. Isolates from planned treatment groups showed an increase in the level of resistance; the lethal concentrations of TBZ to 50% of eggs (LC₅₀s) were 3, 3 and 6 times the LC₅₀s of isolates from nil treatment groups for years 3, 4 and 5 respectively. The LC₅₀s of isolates from regular treatment groups were 14 times higher than those from nil treatment groups in each year. To assess the potential for an increase in level of resistance, additional egg assays were done 14 days after treatment with 44 mg kg^t?1 of TBZ on sheep infected with the planned group isolates for each year. This treatment raised the LC₅₀S for years 3, 4 and 5 respectively by 3, 2 and 1–5 times the LC₅₀s of the same isolates which had not been exposed to additional TBZ treatment. The controlled anthelmintic efficiency assay using 44 mg kg^t?1 of TBZ produced a significant reduction in the number of adult and immature worms from the nil isolate but failed significantly to reduce the number of worms from the planned and regular isolates. A three component analysis resolved the nonlinear trends of the log dose-probit plots in egg hatch assays for isolates from planned treatment groups into subpopulations of susceptible, hybrid and resistant individuals each with different levels of resistance. The proportions of these subpopulations changed in accordance with the level of resistance observed in each year.     

Closure of the micropyle during embryonic development of some pelagic fish eggs

The fine structure of the egg envelope and micropyle was studied in unfertilized and developing eggs of the flounder Paralichthys olivaceus (Temminck & Schlegel), the Alaska pollack Theragra chalcogramma (Pallas), the Japanese tilefish Branchiostegus japonicus (Houttuyn) and the porgy Pagrus major (Temminck & Schlegel). The outer envelope surface of the unfertilized egg was wrinkled, while the inner surface was folded. The micropyle of the unfertilized egg consisted of a shallow vestibule and a distinct canal. The micropylar region of the inner surface of the envelope had a conical- or bowl-shaped protrusion. In developing eggs, the thickness of the envelope decreased and showed smooth outer and inner surfaces which indicated that it had been stretched tangentially at the time of the perivitelline space formation. The lumen of the micropylar canal was invariably occupied with envelope material. We postulate that the blockage of the micropylar canal is a result of the stretching of the envelope. The closure of the micropyle inhibits sperm and external pathogens from penetrating into the perivitelline space and seems to be involved in both the permanent prevention of polyspermy and the protection of the developing embryo from bacterial infection.