Adaptive and evolutionary significance of a reproductive thermal threshold in Barbus barbus

From 1989 to 1996, barbel in the River Ourthe started spawning under variable environmental conditions, except for water temperature. Each year, spawning was initiated when water daily minimum temperature reached or exceeded 13·5° C. Any decrease of temperature below this value later in the spawning period caused spawning to be suspended. Analyses of offspring growth provided evidence that 13·5° C was the value below which 0+ barbel stop growing. It was hypothesized that barbel trade off the lower initial probability of survival against a larger size at the onset of winter. To test empirically for this hypothesis, the adequacy of alternative—theoretical—strategies associated with other thermal thresholds (12, 15·0, 17·1 and 20·2° C) was modelled with respect to: (1) the feasibility of spawning (inhibition of sexual maturation by a decreasing photoperiod); (2) the impact of the temperature on embryonic development; (3) the effect of water level variations on the integrity of spawning grounds until the emergence of larvae; (4) the size of the offspring at the onset of winter. On an 8-year (1989–1996) average, the present spawning strategy would have produced a higher recruitment than alternative strategies (relative adequacy of 33·23, 85·64, 93·17 and 17·62%, respectively). However, alternative strategies would have produced better annual scores on five of eight occasions in the River Ourthe environment, and a better overall score in environments 1·5 or 3·5° C warmer than now. The consistency of the thermal threshold over years, despite a low selection pressure by the environment, was interpreted as the expression of a phenotypic mechanism (thermal homing) promoting the selection of the lowest efficient thermal threshold, and enabling breeders to relay to the next generation some form of thermal stability in a variable environment.     

Reproductive migration of brown trout in a small Norwegian river studied by telemetry

The movement of 34 large (39–73 cm standard length) brown trout Salmo trutta was monitored using radio telemetry for up to 74 days in Brumunda, a small Norwegian river (mean annual discharge 3·3 m³ s⁻¹) flowing into the large Lake Mjøsa. The maximum range of movement in the river was 20 km. No clear relationships existed between individual movement and water discharge, temperature and barometric pressure. Brown trout migrated at all levels of water discharge. At low discharge (<2 m³ s⁻¹) movements were nocturnal. A weir 5·3 km from the outlet restricted ascending brown trout at low ( c . 6° C), but not at high ( c . 8° C) water temperatures. Spawning occurred in September to October and tagged individuals spent 2–51 days at the spawning sites. Mean migration speed from tagging to when the fish reached the spawning area, and from when they left the spawning areas and reached the lake was 1·0 and 2·3 km day⁻¹, respectively. All tagged brown trout that survived spawning returned to the lake after spawning.     

Plasma non-esterified fatty acid profiles in wild Atlantic salmon during their freshwater migration and spawning

Total plasma non-esterified fatty acid (NEFA) levels increased significantly in adult Atlantic salmon during the first months of their upstream migration and spawning in the Exploits River, Newfoundland, Canada. The highest levels occurred in May and were 5467±270·43 nmol ml⁻¹ for females and 4617±334·70 nmol ml⁻¹ for males. Significantly higher levels were maintained by females compared with males for most of the upstream migration. Between August and October, total plasma NEFA levels declined by 61% in females but only 23% in males. The decline in plasma monounsaturated and polyunsaturated fatty acid levels accounted for 74% of the loss of NEFAs in females. Specific plasma NEFAs such as 16: 0 (palmitic), 16: 1 (palmitoleic), 18: 1n9 (oleic) and 20: 5n3 (eicosapentaenoic acid) differed significantly between males and females during migration and spawning. The mean gonadosomatic index ( I _G) values of females in May and just prior to spawning were 0·37±0·01 and 10·25±0·32, respectively. The rapid decline in the plasma NEFA content of females coincided with the largest increase in their I _G (1·85±0·02–10·25±0·32). Corresponding I _G values for males were 0·34±0·01 in May and 3·33±0·78 prior to spawning. Plasma NEFA levels of spent salmon did not differ between sexes and were significantly lower than those of salmon preparing to spawn.     

Seasonal Timing and Diel Activity of Lacustrine Brook Charr, Salvelinus Fontinalis, Spawning in a Lake Outlet

The objectives of this study were to determine (1) the seasonal timing and diel activity patterns of lacustrine brook charr when migrating to a lake outlet for spawning, and (2) the homing of reproducing individuals. In 1995, 745 individuals were captured while migrating to the spawning ground compared to 1148 in 1996. In both years, the sex ratio was not significantly different from 1:1 but females were significantly longer than males. The spawning migrations lasted about 48d and were more nocturnal (between 20:00 and 8:00h) than diurnal. The migration to the spawning ground began when water temperature decreased to 13°C and peaked when water temperature decreased from 11°C to 7°C. Peaks in migration activity were always preceded by a sudden decrease followed by an increase in temperature. The spawning migration lasted until the third week of October, when water temperature varied between 3°C and 8°C. Males, especially larger ones, arrived on the spawning ground before females in both years. The length of males migrating to the spawning ground decreased during the spawning season while the females' length showed no pattern. Only 9.7% of the 745 fin-clipped individuals from 1995 returned to the lake outlet to spawn in 1996. The results of this study indicate that lacustrine brook charr show similarities to other anadromous and lake-spawning salmonid populations when migrating to spawning grounds.     

Reproductive strategy and spawning activity of the European hake Merluccius merluccius (L.) in the Bay of Biscay

Histological examination of ovaries from European hake Merluccius merluccius from the Bay of Biscay sampled on a monthly basis from December 1996 to October 1997 demonstrated that European hake exhibit indeterminate fecundity. Oocyte development is asynchronous with a continuous oocyte size-frequency distribution in prespawning, spawning and postspawning females, and a gradual decrease in the mean diameter of advanced yolked oocytes as spawning proceeds. Spawning European hake were observed throughout the 11 month study period. The spawning fraction ranged from 0·085 to 0·207, which is equivalent to a batch interval of 5–12 days. The spawning fraction and proportion of spawning females were highest from January to March. Seasonal variation of atresia was inversely related to spawning fraction. Extensive atresia in individual fish was observed in September and October, when the spawning fraction was low.     

Climate variables and spawning migrations of the striped mullet and white mullet in the north‐western area of the Gulf of Mexico

The 'smoothed'(4253e HT smoother) 6 year time series with a monthly periodicity of catches (where the increases of the catches reflect the spawning migration) of Mugil cephalus and Mugil curema , were correlated (Pearson correlation index) against the 'smoothed' data on rainfall, water temperature, daylight and tide monthly averages. Water temperature and M. curema spawning migrations had a negative correlation ( r  = −0·77), while tide monthly averages and M. cephalus spawning migrations were positively correlated ( r  = 0·84). The two environmental variables that showed highly significant correlations were water temperature and daylight ( r  = 0·85, P  < 0·05). Meteorological tides were used as a reflection of the effect of the north winds from the Gulf of Mexico which produce an average tide rise from September to November. The relationship of these variables with spawning migrations is discussed, and it is suggested, in particular, that meteorological tides, as a reflection of the north winds, are the trigger (zeitgebers) for M. cephalus spawning in the ocean.     

Characteristics of egg and larval production in captive bluespotted gobies

Spawning of the Hawaiian coral-reef goby Asterropteryx semipunctata was diurnal, occurring at various times throughout the day. Mean length of eggs deposited in nests was 0·76 mm (range 0·67–0·84); mean egg width was 0·47 mm (range 0·41–0·52). Clutch size varied from 296 to 1552 eggs (mean=886±309), and was independent of standard length, total body weight, and body condition. Mean relative clutch size was 1·59 eggs mg^-1 total body weight (range 0·84–2·43). Clutches hatched 4–5 nights after being deposited in a nest. Mean notochord length of newly-hatched larvae was 1·88 mm (range 1·60–2·04). The minimum period of time that elapsed between egg deposition and subsequent growth of a new batch of oocytes to spawning size was 5–6 days, providing a reasonable estimate of minimum spawning interval. Compared with other gobiids, tropical species tend to have shorter incubation periods, smaller eggs and smaller larvae at hatching.     

Spawning characteristics of the anadromous brown trout in a small Swedish stream

The spawning pattern of the anadromous brown trout Salmo trutta was studied in Själsöån, a small stream in Gotland, Sweden, during eight winters between 1992–1993 and 1999–2000. The total length ( L _T) at spawning was normally distributed (185–890 mm) for females and multimodal for males (300, 400 and 550 mm most frequent length classes). Spawning males were significantly younger (2+ to 4+ years) than females (3+ to 5+ years). The sex‐ratio at the beginning and at the end of the spawning season favoured males. The mean ±  s . d . number of spawners was 70 ± 16 individuals per year. Migration into and out of the stream occurred between November and June. The highest number of spawning fish was found in the stream at the end of November or at the beginning of December. Migration mainly occurred during high water flow and at night. The majority of the females entered the stream and spawned the same (29·3% of all the females) or the next night (32·8% of all the females) while males may have stayed for 2 to 3 weeks (21·3% of all the males) in the stream before spawning. Males usually remained much longer in the stream (mean ±  s . d . 45 ± 56 days) than females (16 ± 30 days). Females lost more mass in the stream (mean ±  s . d . 17·3 ± 8·6%) than males (7·7 ± 9·6%). For both sexes, mass loss was positively correlated with the time spent in the stream. Only 7·3% of the males and 5·7% of the females occurred in the stream for >1 year. Spawning took place only during the night.     

Reproductive behaviour of a temperate serranid fish, Paralabrax clathratus(Girard), from Santa Catalina Island, California, U.S.A.

The reproductive behaviour of the kelp bass Paralabrax clathratus was studied on Santa Catalina Island, California, U.S.A. from April 2000 to September 2002. Adults formed aggregations of three to > 200 individuals, and spawning occurred within subgroups of three to 23 individuals that contained a single female. The gonado‐somatic index ( I _G) of collected ripe males (mean = 5·8%, range = 0·5–13·1%) indicated a large investment in sperm production that is common in group‐spawning fishes characterized by intense sperm competition. Spawning occurred 32 min before sunset to 120 min after sunset, and both males and females were capable of spawning multiple times during a single evening. Behavioural observations of adults and estimates of spawning periodicity from the collection of females with hydrated oocytes suggested that spawning occurred continuously throughout the summer months and showed no significant relationship with the lunar cycle. In general, the spawning behaviour of kelp bass was similar to other functionally gonochoric, group‐spawning serranids. The dynamics of P. clathratus spawning aggregations, however, were inconsistent with that of tropical reef fish spawning aggregations, including the transient spawning aggregations of some tropical serranids. Aggregation spawning appeared to be an important component of the annual reproduction of this species.     

Reproductive biology of the endangered Oxleyan pygmy perch Nannoperca oxleyana Whitley

The reproductive biology of the Oxleyan pygmy perch Nannoperca oxleyana is described from simultaneous studies of wild populations in north-eastern New South Wales and mature fish held in aquaria. In the wild, 50% of males and females matured at total lengths of 24·0–25·9 and 28·0–29·9 mm, respectively. The species displays sexual dichromatism during the spawning season, with males developing more intense red and brown fin and body colouration, and black pelvic fins. Captive male broodfish displayed territoriality during the breeding season, closely guarding sites within artificial, plant-like substrata in which pairs of fish spawned adhesive eggs. Protracted serial spawning of wild and captive fish occurred from September to April and May at mean water temperatures ≥16·6° C and day length ≥10·7 h. Captive broodfish spawned on an average of 57% of days during the 256 day spawning period. Gonado-somatic indices averaged 0·7% for all ripe males and 4·1–4·2% for all ripe females collected. Mean total and batch fecundities of captive females were 1323 eggs per fish and 7·8 eggs per fish per day, respectively, and relative fecundity was 587 eggs g⁻¹ of body mass. Batch fecundity of wild females was estimated at 7·8 eggs per fish. The adaptive significance of this reproductive strategy in a harsh, variable environment is discussed.     

Reproductive biology of Vinciguerria nimbaria in the equatorial waters of the eastern Atlantic Ocean

Adult Vinciguerria nimbaria in an area of the Atlantic Ocean (0–5°N; 10–20°W), collected from November 1994 to February 1998, exhibited two different patterns in diel behaviour: 'typical' behaviour which consisted of large diel vertical migration and 'atypical', characterized by concentrations of schools that remained in the surface layers during the daytime. The total life span of V. nimbaria was estimated at 6–7 months. Females were mature when they reached 30·6 mm standard length ( L _S), which corresponded to an age of 85 days. Once the females were mature, spawning was continuous in the population as a whole, and V. nimbaria spawned continuously throughout the year. Spawning took place during two restricted times of the day: typical females spawned at dusk and atypical females spawned at dawn. Using the post-ovulatory follicles method, spawning frequency was estimated at 2 days. Batch fecundity was estimated at 1236 oocytes or 1230 oocytes g⁻¹ of total body mass, and egg size was 650 µm whatever the period of the year. The lifetime fecundity of V. nimbaria was estimated at 9000 eggs (109 000 eggs if mortality rate was not taken into account), and the maximum stock egg production of a theoretical cohort occurred at 37 mm L _S. Young adults thus contributed the most to the reproductive output for the survival of the population.     

Reproductive biology of gag in the southern Gulf of Mexico

Aspects of the reproductive biology of gag Mycteroperca microlepis in the southern Gulf of Mexico were studied by following seasonal variations in the gonado‐somatic index and through histological examination of gonads. Gag were collected from inshore and offshore waters of the Campeche Bank, Yucatan, Mexico, between April 1996 and December 2001. This species is a protogynous hermaphrodite, and appeared to be depth‐size distributed. The smallest gag (9–49 cm L _F) collected were all juvenile females, and were caught in inshore waters (1–10 m depth), while the largest (49–116 cm L _F), mainly adult females, males and transitionals, were captured in offshore waters (33–167 m depth). Overall the offshore male to female ratio was female‐biased (1 : 3·3) and differed significantly from unity. The species spawns at depths of c . 50–53 m, from early winter to mid‐spring, with peak spawning activity occurring between January and March. Fifty per cent of females reached first maturity at 72·1 cm L _F. At 103 cm L _F, 50% of sampled females had changed into males. Gag can be considered a monandric species, and sexual transition for this grouper seemed to occur in fish distributed within a narrow size range (85–111 cm L _F). The results are compared with those of other authors for gag stocks from the south‐east Atlantic coast of the U.S.A. and the north‐east Gulf of Mexico.     

A preliminary investigation of spawning migrations of grayling in a small stream as determined by radio-tracking

Adult grayling Thymallus thymallus migrated from 230 to 4980 m up the Aisne stream, Belgium, to spawn between 18 and 29 March, under decreasing floods, increasing temperature and low turbidity. Males ( n =4) arrived on spawning grounds several days earlier than females ( n =2), stayed there longer (10–19 v. 2–3 days), and occupied a single ground each, whereas females moved between several places. After spawning, all grayling homed precisely into the pool-riffle sequences where they were tagged in late February, and remained here until late June. These observations indicate that resident grayling are far less mobile than autumn-spawning salmonids, and that the environmental factors triggering spawning migrations resemble more closely those of spring-spawning cyprinids than of other salmonids. The implications of these restricted mobility patterns are discussed within the scope of population structure, and impact of river management.     

The reproduction, age and growth of the spotted rockling

Biology of the Macaronesian endemic rockling Gaidropsarus guttatus was studied in the Azores. The overall sex ratio from the samples was highly in favour of females (1 : 6·33). The growth parameters were L _∞ = 24·23, k  = 1·219 and t ₀ = −0·059. Fish matured at 15 cm L _T and the spawning season was strongly clustered in April.     

Reproductive biology and recruitment of the white sea bream in the Azores

The life history of the white sea bream Diplodus sargus in the Azores showed a pattern consistent with digynic hermaphroditism achieving sexual maturity during the second year of life, at 16·7 cm L _T. Spawning occurred from March to June at temperatures between 15 and 17° C and the onset and duration of spawning season in the sea bream appeared to be influenced by sea water temperatures. As latitude decreased, both in the northern and southern hemispheres, the spawning season of D. sargus populations started earlier and extended longer, highlighting the potential importance of temperature to the onset and duration of reproduction in this species. Settlement took place from late May to July, and settlers remained in the nursery area for c . 2·5 months. Emigration from the nursery area to join shoals of juveniles occurred from late July to September.     

Reproductive ecology of the river lamprey

The reproductive ecology of river lamprey Lampetra fluviatilis was investigated during the spawning period 2003 in the River Derwent, north‐east England. Over this period 1199 ± 104 individuals day⁻¹(mean ±  s . d .) were counted on one spawning site (area c . 450 m²), but mark‐recapture estimates suggested that >5000 river lamprey used this site over the same period and egg deposition was estimated as 168 000 eggs m⁻². The operational sex ratio of river lamprey in spawning clusters changed between spawning phases, from domination by females during the nest‐building phase (male : female ratio, 1 : 3·46), to a preponderance of males during the spawning phase (male : female, 1 : 0·37), followed by a return to a majority of females after spawning (male : female, 1 : 3·74). Recapture data showed that >97% of recaptured, tagged males were recorded at two or more nests, whereas almost 50% of recaptured, tagged females were recorded at the same nest, suggesting a promiscuous mating system, with a tendency towards polygyny within the population. Within the lower 80 km of the River Derwent and its tributaries, evidence of river lamprey spawning was found at only six sites, and most spawning (>80% of the observed spawning population) was at one site.     

Sex structure and potential female fecundity in a Epinephelus guttatus spawning aggregation: applying ultrasonic imaging

Ultrasonic imaging was used to determine the spawning population structure and develop a fecundity estimation model for a red hind Epinephelus guttatus spawning aggregation within the Red Hind Bank Marine Conservation District, St Thomas, U.S.V.I. The spawning population showed considerable within‐month and between‐month variation in population size‐ and sex‐structure. In the spawning season studied, males appeared to arrive at the aggregation site first in December although females represented a large proportion of the catch early in the aggregation periods in January and February. Spawning occurred in January and February, and size frequency distributions suggested that an influx of small females occurred during the second spawning month. An overall sex ratio of 2·9 : 1 (female : male) was recorded for the whole reproductive season. The sex ratio, however, differed between months and days within months. More females per male were recorded in January than in February when the sex ratio was male biased. Fecundity estimates for this species predicted very high potential fecundities (2·4 × 10⁵−2·4 × 10⁶ oocytes). The ultrasound model also illustrated a rapid increase in potential female fecundity with total length. Ultrasonic imaging may prove a valuable tool in population assessment for many species and locations in which invasive fishing methods are clearly undesirable.     

Reproductive biology, species status and taxonomic relationships of four recently discovered galaxiid fishes in a New Zealand river

The reproductive biology of four recently discovered, largely allopatric non-migratory Galaxias species in the Taieri River, New Zealand, was compared with the closely related Galaxias vulgaris . Timing of spawning was similar (late winter/spring) but spawning habitat, spawning behaviour and nest structure differed among the Taieri species and G. vulgaris . Patterns of fecundity also differed among the Taieri species but, with G. vulgaris , they form a distinct group of galaxiids with maxima of 800–1500 eggs having diameters of 2·3–2·8 μn. The different spawning behaviours of the Taieri galaxiids, in particular the differences in spawning orientation, provide evidence for prezygotic isolating mechanisms and support their status as separate species. In cladistics terms, the results indicate that Galaxias anomalus, G. eldoni and G. vulgaris are sister taxa, while G. depressiceps and Galaxias sp. form a second cluster.     

Reproductive biology of the threatened golden galaxias Galaxias auratus Johnston and the influence of lake hydrology

Golden galaxias Galaxias auratus (31–235 mm fork length, L _F) were collected monthly from littoral habitats in Lakes Crescent and Sorell, Tasmania, Australia, between July 2000 and December 2002. Spawning habitats were identified and monitored in both lakes, and surveyed in Lake Crescent. Trends in gonado-somatic indices and reproductive stages of development indicated that gonad development in both sexes begins in midsummer and peaks in late autumn to early winter. Males mature at smaller sizes (50% at 52 mm L _F) than females (50% at 76 mm L _F), larger individuals are predominately females (95% of fish ≥138 mm L _F), and overall male to female ratios are female biased ( c . 1:2). Spawning occurs late autumn to early spring (water temperatures = 1·4–9·7° C) with peaks in spawning activity in winter (mean water temperatures <5° C). Demersal adhesive eggs ( c. 1·5 mm diameter) were found on cobble substrata ( c. 20–250 mm diameter) in littoral areas ( c. 0·2–0·6 m deep) and fecundity of fish 71–181 mm L _F ranged from 619 to 14 478 eggs. The rate of change in water level over the 20 days prior to monthly sampling was important in explaining the occurrence of spent fish and this accounted for temporal differences in spawning between the populations. Lake hydrology influences the reproductive cycle of G. auratus by possibly providing a stimulus for spawning and it controls the availability of spawning habitat in Lake Crescent. Seasonal hydrological cycles ( i.e. rises during late autumn to winter) and a minimum water level of 802·20 m Australian Height Datum in Lake Crescent during autumn (above which littoral areas of cobble substratum are inundated) are critical to G. auratus populations.     

The role of life history in the relationship between population dynamics and environmental variability in two Mediterranean stream fishes

Chub Squalius torgalensis and nase Chondrostoma lusitanicum , in a Mediterranean stream, showed important differences in life-history traits and population dynamics. Both species reached mean maturity at age 2 years. Chub lived up to age 5 years, spawned in March to June, grew at a maximum rate of 0·59 mm mm⁻¹ year⁻¹ and showed a low reproductive allocation, with fecundity and egg size increasing with body size. Nase lived up to age 4 years, spawned in January to April, grew at a maximum rate of 0·46 mm mm⁻¹ year⁻¹ and showed a high reproductive allocation, with egg size independent of body size. Both chub and nase showed moderate fluctuations in population size during 1991–1998, but differed in factors driving density at age. Density of age 1 year juvenile chub decreased following severe summer droughts and proportionate survival prevailed thereafter. Density of age 2 year adult nase decreased following severe spring floods, but neither environmental nor parental stock effects were detected for juveniles and older fishes. The results illustrated the interplay between life history and environmental variability in driving fish population dynamics, with impacts of both summer droughts and spring floods being contingent on species-specific patterns of spawning and reproductive investment.