Reproductive biology ofAwaous guamensis, an amphidromous Hawaiian goby

Synopsis Spawning season, size at first reproduction, oocyte maturation, and fecundity ofAwaous guamensis, an amphidromous Hawaiian goby, were studied from June 1989 through May 1991 in the Wainiha River, Kau'ai, Hawai'i. Female fish larger than 73 mm standard length (SL) had mature gonads from August through December in 1989 and 1990. Gonadosomatic index (GSI) values for mature females ranged from 0.2 to 14.5 during the spawning season. Male fish larger than 64 mm SL had elevated GSI values from June 1989 through December 1989 and from August 1990 through December 1990. Mature sperm were found in two male fish collected in January and February. GSI values for mature males ranged from less than 0.01 to 4.0 in the spawning season. Size-frequency distributions of measurements of vitellogenic oocyte diameters and microscopic observations of oocytes indicated this species has group-synchronous oocyte development. Ovarian maturation stages examined over a 29-month period suggest that members of the stock spawned at different times within the spawning season, although mass spawning events have been documented for this species. Estimates of clutch sizes from nests measured in situ were comparable to estimates of potential fecundity from in vitro examination of ovaries, and indicated that female fish deposited an entire clutch during a spawning event. No evidence for multiple spawning by an individual fish in a single season was found. However, microscopic observations of brown bodies in some ovaries suggested that individual fish probably spawn more than once in a lifetime.     

Intra-specific variations of egg size,clutch size and larval survival relatedto maternal size in amphidromous <Emphasis Type="BoldItalic">Rhinogobius</Emphasis> goby

Synopsis Newly hatched amphidromous Rhinogobius sp. CB (cross band type) larvae drift downstream to the sea to grow and develop before returning upstream as juveniles. Since larger and older individuals usually inhabit the upper reaches of rivers, larvae from larger females are more likely to suffer higher risks of starvation or predation during their longer migration to the sea. We examined the relationship between reproductive parameters (egg volume and clutch size) and maternal length. We collected adult Rhinogobius sp. CB along the course of the Aizu River (Wakayama Prefecture, Japan) and spawned them under laboratory conditions. We measured egg volumes and clutch sizes, as well as larval starvation tolerance. Both egg volume and clutch size increased with standard length or age of the maternal fish, while egg density in these clusters did not correlate with standard length. Gonad-somatic index (GSI) also tended to increase with maternal standard length. There was a significant positive correlation between egg size and the 72-h survival rate of unfed hatchlings. Intra-specific variation of egg volumes and clutch sizes in this species seems to be an adaptation for enhancing offspring survivorship during migration to the sea. Some females spawned a second time. Second spawned egg sizes were smaller than first spawned egg sizes, although there was no difference in clutch size between the two. Egg size variation between subsequent spawns may be an adjustment to the changes in seasonal environmental conditions.     

Variation in male mating success in the paternal nest brooder, Rhinogobius goby, along a river course

Rhinogobius sp. CB (cross band type) is an amphidromous freshwater goby which receives paternal nest care under stones on river beds. This goby is known to migrate to the upper reaches of rivers as it grows. In the present study, the relationships among male mating success of Rhinogobius sp. CB, male body size, the size frequency distribution of river bed stones and the presence of the sympatric goby Tridentiger brevispinis (which uses nest sites similar to those of Rhinogobius sp. CB) were investigated along the course of the Aizu River, Wakayama Prefecture, Japan. In the lower reach, where T. brevispinis was present and large stones were scarce, the sizes of the nest stones and the egg clusters of male Rhinogobius sp. CB were smaller than those of T. brevispinis. In the middle and the upper reaches, where T. brevispinis was absent and large stones were abundant, males of Rhinogobius sp. CB used larger nest stones than in the lower reach. In both gobies, there were positive correlations among male body size, nest size and egg cluster size in all reaches. Nest-choice experiments carried out in the laboratory, with or without a rival, showed that males of both gobies preferentially used large nest sites and that large males had an advantage in male-male competition for large nest sites. These results indicate that large male body size can increase male mating success and that male mating success increases in the upper reach in this goby. The migration pattern of Rhinogobius sp. CB was also discussed in relation to these findings.     

Variations in clutch and egg sizes in the amphidromous goby <Emphasis Type="Italic">Rhinogobius</Emphasis> sp. CB along a river course and within a spawning season

Rhinogobius sp. CB is a common amphidromous goby in Japan. I investigated egg and clutch sizes along a river course and within a spawning season in the Aizu River, Kii Peninsula, Japan. Clutch size within the same female size did not differ along the course of the river, but decreased from early to later in the spawning season. Egg size within the same female size decreased from early to later in the spawning season and from the upper to the lower reaches. These results mean energy resources allocated to the ovary decreased seasonally and locally. Several factors are presented to explain these variations, such as female somatic conditions, a tradeoff between reproduction and growth, food availability, water temperature, and presence of competitive species. Some factors explaining the seasonal and local variations in egg size, such as water temperature and food availability for offspring, are also presented. Egg size correlated well with female size. The trend in egg size variation with female size seems to be an adaptation for their migrating pattern, as larvae from larger female, which tended to inhabit the upper reaches, would have a greater endurance against starvation during their migration to the sea.     

Distribution of hydroids along fronds of the kelp Ecklonia radiata

A full understanding of life history characteristics of invasive species is a fundamental prerequisite for the development of management strategies. Two introduced goby species (Rhinogobius cliffordpopei and Rhinogobius giurinus) have established highly abundant populations in Lake Erhai (China). In the present study, we examined the reproductive biology of these two species with the aim of improving the efficiency of management strategy. The results indicated that R. cliffordpopei spawned from February to June, whereas R. giurinus spawned from April to August. Rhinogobius cliffordpopei showed higher gonado-somatic indices and had larger eggs than R. giurinus. The adult sex ratio of R. cliffordpopei was female skewed, but that of R. giurinus was equal. Rhinogobius cliffordpopei showed a male-skewed sexual size dimorphism, whereas the body size of R. giurinus varied only slightly between males and females. The different reproductive traits appear as a crucial biologic aspect for developing control programs. Specifically, control measures should be implemented and/or intensified from September to February for R. cliffordpopei and from January to April for R. giurinus. The body size of the smaller R. cliffordpopei females is the determinant for minimal mesh size of the nets used in physical removals of R. cliffordpopei.     

Reproductive biology of an amphidromous goby, Sicyopterus lagocephalus, in La Réunion Island

In the Indo-Pacific region, rivers are inhabited by amphidromous gobies. They are often subjected to a heavy fishing pressure, highlighting the urgent need to acquire knowledge on their biology for management purposes. This study investigated the reproductive strategy of Sicyopterus lagocephalus, a widespread amphidromous goby, in two rivers of La Réunion Island. Histological observations of ovarian samples and oocyte-size frequency distributions revealed that females had a group-synchronous ovarian follicle development (i.e., two cohorts of oocytes were distinguished simultaneously in ovaries). Females laid an entire clutch in a unique event (from 14,304 to 232,475 eggs) and then another batch of oocytes was recruited, showing that the annual fecundity is indeterminate. Ovarian growth was isometric at all oocyte development stages showing that the gonadosomatic index (GSI) is a good proxy of reproductive condition. The main reproductive season of S. lagocephalus spanned from early February to May with a narrower range upstream and a wider one downstream. Reproduction activity is mainly restricted by water temperatures in upstream areas, whereas it is likely influenced by female body condition and competition in downstream areas. The variability of reproductive traits leads to size structure variation in stocks of spawning females throughout the year.     

The reproductive ecology of two subspecies of the bitterling Rhodeus atremius (Cyprinidae, Acheilognathinae)

The reproductive ecology of two endangered subspecies of the bitterling Rhodeus atremius was investigated in two rivers in Okayama and Fukuoka Prefectures, Japan. Several other bitterling species, R. ocellatus, Tanakia limbata, T. lanceolata, Acheilognathus tabira and A. rhombeus, are also found in these rivers. Spawning of both R. atremius subspecies was recorded between late March and August 2007, peaking between April and July. Females of R. atremius began to mature at small body size (minimum 25 mm). They had small clutch sizes (maximum 16 eggs) and developed a short ovipositor (mean 13.6 mm), which was used to deposit a bulb-shaped egg of comparatively large size (3.2 mm³, major and minor axes 2.8 and 1.5 mm). Clutch size, ovipositor length and egg size appear to be a function of female body size. Both the body size and ovipositor length of R. atremius with ripe ova and the size and shape of the eggs varied seasonally. Rhodeus a. suigensis achieved maturity at a smaller body size, had a longer ovipositor and produced smaller eggs, with the minor axis being wider relative to the major axis than those of R. a. atremius. Rhodeus atremius may be adapted for depositing small numbers of eggs in a much shallower position inside the gills of smaller host mussels than other bitterling species.     

Life history variation in males of the river sculpin,Cottus hangiongensis,along the course of a river

Synopsis Body size at maturity, individual growth, upstream migration and spatial stability in males of the amphidromous sculpin,Cottus hangiongensis (Cottidae), were investigated in the Daitobetsu River of southern Hokkaido, Japan. The body size of males at first maturity varied gradually along the course of the river, from approximately 70 mm SL in the lowest reaches (0.2 km up from the river mouth) to about 120 mm SL in the uppermost reaches (about 6 km up from the river mouth). The age at first maturity was estimated as 2 to 3 years in the lower reaches and as 4 to 5 years in the upper reaches. Individual growth rate tended to increase upstream, while population density decreased. These results suggest that the male life-history pathway varies along the course of the river and so allows one to consider the relationships to the polygynous mating system of this species in terms of current hypotheses regarding alternative reproductive tactics.     

Reproduction and early development of a freshwater pipefish <Emphasis Type="Italic">Microphis leiaspis</Emphasis> in Okinawa-jima Island,Japan

We investigated the size at maturation, breeding season, and morphological development of larvae and juveniles of a freshwater pipefish Microphis leiaspis, which belongs to Gastrophori, collected from three rivers on the northern part of Okinawa-jima Island, Japan. The minimum size of brooding males was 105–123 mm in standard length (SL). The smallest mature female was estimated to be ca. 130 mm SL from the analysis of gonadosomatic index (GSI) and histological observations of gonads. The breeding season was estimated to be from June to December according to monthly changes in female GSI, histological observations of gonads, and monthly changes in the occurrence of brooding males. The number of eggs in the male brood pouch ranged from 75 to 241 (mean ± SD: 152 ± 52, n = 22). The male releases newly hatched larvae in freshwater areas. After newborns grow in the sea, they return to freshwater areas of the rivers and attain maturity. Microphis leiaspis was conformed to have an amphidromous life history. Notochord length of the released larvae was 6.1 mm, with a well-developed finfold. Larvae attained 11.1 mm SL, formation of the caudal and dorsal fin rays was complete, and the caudal fin became lozenge shaped at 30 days after the release, and juveniles reached 36.0 mm SL at 63 days after release. In the period between 30 and 63 days after the release, formation of all fins except the pectoral fins was completed, and caudal fin rays were extended and sector shaped with deep slits between each fin ray. The morphology of the released larvae of M. leiaspis is similar to that of Gastrophori species, and the morphology of juveniles similar to other species of Microphis.     

Seasonal variation of egg size and number in a Daphnia pulex population

Seasonal variation of egg size and number was examined in a Daphnia pulex population inhabiting a vernal pond. In this population, size at maturity declines at midseason, probably as an adaptive response to size-selective predation by larvae of the salamander Ambystoma. The larger early season individuals produce more and larger eggs than the smaller late season individuals. Age at maturity does not vary between seasons. Laboratory experiments indicate that temperature may affect egg size, egg number and size at maturity. However, field data suggest that temperature accounts for only a small fraction of the total variation in egg size and number. Indirect measures of nutrition indicate that food limitation does not cause the seasonal decline in egg size and number. The seasonal change in reproductive traits is well correlated with changes in invertebrate and vertebrate predation. Examination of predator feeding preferences and their impact on Daphnia mortality indicate that variation of reproductive traits is most likely a complex adaptation to changing predation regimes.     

Reproductive potential of Daphnia galeata in relation to food conditions: implications of a changing size-structure of the population

A field study was conducted in the highly eutrophic, biomanipulated Bautzen Reservoir to investigate the influences of food conditions on fecundity of Daphnia galeata during spring and early summer in four consecutive years. By defining a relation between body size and maximum brood size, the realised reproductive potential (RRP) of Daphnia was expressed as percentage of actual egg numbers of the population in relation to calculated maximum egg numbers. This parameter is independent of the size structure of the population. It was found to correlate generally with mean clutch size. However, deviations occurred if size at maturity (SAM) declined. Extremely low SAM values (0.8 mm) were recorded in 2 years, corresponding with increasing food concentrations. Low clutch sizes in these situations can be explained by small body sizes of ovigerous females, whereas RRP increased, thus reflecting good food conditions. From the comparison of different reproductive parameters, it is concluded that RRP is the most reliable one with regard to bottom-up influences in case of extremely variable size structure and possible predation impacts. Fecundity of D. galeata was clearly food limited during the clear-water stage. Relatively high POC-threshold concentrations of 0.26 mg C l^–1 suggest that food quality was low and may be equally important as food quantity in limiting egg production.     

Simultaneous maturation of two discrete oocyte batches in ayu

Ayu Plecoglossus altivelis including amphidromous and landlocked populations, are known for batch spawning. Unusual clutches at ovulation were observed from rearing stocks that were naturally induced to sexual maturation. Two peaks occurred simultaneously in the frequency distribution of egg size, compared to normal fish, which have clutches with unimodal size distribution. The eggs in the small size class could be fertilized, and small-sized larvae hatched from these eggs. To explain the alteration in egg size through divergence from amphidromous to landlocked populations, the role of batch allocation among oocytes is discussed.     

Reproductive biology of an endemic,amphidromous goby Lentipes concolor in Hawaiian streams

Synopsis Egg size, fecundity, nest site selection, and breeding structure of the amphidromous Hawaiian goby Lentipes concolor were studied to determine if there was any relationship between these aspects of the breeding biology of this fish and the very long larval period of Hawaiian stream gobies. To do this, these parameters were compared with those known for other gobioid fishes. While eggs were small and females were found to lay up to four nests in a season, reproductive biology and mating behavior of L. concolor were typical of many gobies in spite of its amphidromous life history pattern. Lentipes concolor nests were found from October to June with a spawning peak in February. Nests tended to be positioned closer to the stream bank, and under larger rocks than if they were situated randomly. Depth and mean water column velocity over nest sites did not differ from that at randomly selected points in the stream.     

Fecundity and oviposition ofEucelatoria bryani, a gregarious parasitoid ofHelicoverpa zea andHeliothis virescens

We examined longevity, fecundity, and oviposition strategies ofEucelatoria bryani Sabrosky (Diptera: Tachinidae), a gregarious endoparasitoid ofHelicoverpa zea (Boddie) andHeliothis virescens (F.) (Lepidoptera: Noctuidae). Longevity of adult femaleE. bryani was not related to body size. In contrast to longevity, largerE. bryani females had greater potential fecundity than smaller females, as determined by the number of embryonated eggs present in the common oviduct. However, female parasitoid size did not affect primary clutch size (number of eggs deposited in a host). Because embryos in eggs located in the ovisac were larger than those located elsewhere in the common oviduct, maximum primary clutch size may be physiologically limited by the number of fully mature eggs a female has available at one time.E. bryani females adjusted primary clutch size in response to host size, for bothH. zea andH. virescens. This adjustment appears to be adaptive because females did not overexploit hosts by depositing more larvae than a host could support. Adult emergence was not related to host size. Although host weight positively influencedE. bryani progeny weight, increases in progeny size with host size were counterbalanced by increases in primary clutch size with host size.     

Otolith microchemistry of tropical diadromous fishes: spatial and migratory dynamics

Otolith microchemistry was applied to quantify migratory variation and the proportion of native Caribbean stream fishes that undergo full or partial marine migration. Strontium and barium water chemistry in four Puerto Rico, U.S.A., rivers was clearly related to a salinity gradient; however, variation in water barium, and thus fish otoliths, was also dependent on river basin. Strontium was the most accurate index of longitudinal migration in tropical diadromous fish otoliths. Among the four species examined, bigmouth sleeper Gobiomorus dormitor, mountain mullet Agonostomus monticola, sirajo goby Sicydium spp. and river goby Awaous banana, most individuals were fully amphidromous, but 9–12% were semi‐amphidromous as recruits, having never experienced marine or estuarine conditions in early life stages and showing no evidence of marine elemental signatures in their otolith core. Populations of one species, G. dormitor, may have contained a small contingent of semi‐amphidromous adults, migratory individuals that periodically occupied marine or estuarine habitats (4%); however, adult migratory elemental signatures may have been confounded with those related to diet and physiology. These findings indicate the plasticity of migratory strategies of tropical diadromous fishes, which may be more variable than simple categorization might suggest.     

Selectivity Parameters and Size at First Maturity in Deep-Water Shrimps, Aristaeomorpha foliacea (Risso, 1827) and Aristeus antennatus (Risso, 1816), from the North-Western Ionian Sea (Mediterranean Sea)

Selectivity experiments were carried out during trawling targeting deep-water shrimps Aristaeomorpha foliacea (Risso, 1827) and Aristeus antennatus (Risso, 1816) (Crustacea, Decapoda, Aristeidae) in the North-Western Ionian Sea (Eastern-Central Mediterranean). Different criteria were employed to analyse maturity; however, the proportion at 50% of retained, mated and mature specimens was always used to indicate the size, expressed as Carapace Length (CL, mm), at first capture (CL_c), mating (CL_sp) and at first maturity (CL_m), respectively. In order to estimate the size at 50% maturity (CL_m) for females of both species, three criteria were adopted. In particular, CL_m was computed for the mature females not considering the presence of spermatophores, for the mature females with spermatophores and for the mature females intersected by the decreasing proportion with size of females without spermatophores. Three diamond stretched mesh codends of 40, 50 and 60 mm were tested using a cover of 20 mm. The 40-mm stretched mesh size (European Union legal size in the Mediterranean) was not selective for the sampled population of each species. The size at first capture (CL_c), calculated in both species for the two sexes combined, increased significantly with mesh size. Even for the mesh size of 60 mm, the size at first capture was still smaller than the sizes at 50% maturity, whatever the criterion adopted. Since the differences between the size at first maturity and the sizes at first capture are greater in A. foliacea than A. antennatus, the former species appears in this respect to be more vulnerable to trawling than the latter.     

Diet shift response in round goby,Neogobius melanostomus,based on size,sex, depth,and habitat in the western basin of Lake Erie

This study examines the diet of the round goby (Neogobius melanostomus) in the western basin of Lake Erie. As an invasive benthic feeder, the round goby has the potential to affect smallmouth bass and other native species. Round goby (n = 100) were collected during summer 2011 and stomach contents were examined to determine diet patterns and possible ontogenetic diet shifts. Individual round goby were grouped by sex, size (small < 65 mm, large ≥ 65 mm), depth of habitat (shallow < 2 m, deep > 5 m), and habitat type (natural shallows, anthropogenically modified shallows, and deep waters). Gut fullness ranged from 10 to 100% with 23 stomachs presenting 100% fullness. Round goby consumed 27 different food items including abiotic (sand grains, gravel), floral (algae, fine leaved vascular plants), and fauna items. The Index of Relative Importance (IRI) and comparative z‐tests were used to assess correlations and differences. Two‐way z‐tests revealed a significant difference in mouth gape related to size (z = ?5.56377, P = 2.64e‐08), and habitat depth (z = 3.34262, P = 0.00083). A significant linear correlation was also found between mouth gape standardized by head length (HL) for both males (P = 2.63e‐9) and females (P = 1.3e‐4). Two‐way z‐tests also revealed a significant difference in gonadosomatic index (GSI) related to sex (z = 6.07727, P = 6.11248e‐10), but not size. A significant difference in gut fullness was also found related to sex (z = ?3.34743, P = 0.00082), habitat depth (z = 3.16336, P = 0.00156), and habitat type (z = ?2.7398, P = 0.00615). IRI values demonstrated a diet selective of veliger mussels (IRI = 2462.01), juvenile mussels (IRI = 1073.03), cladocerans (IRI = 4804.31), and chironomids (IRI = 1012.12). While previous studies have focused on round goby diet shifts from macroinvertebrates to bivalves, most studies did not evaluate changes in diet among multiple categories. Furthering knowledge of multiple aspects of goby diet may aid in developing management techniques to deter future round goby invasion.     

WHY ARE CLUTCH SIZES MORE VARIABLE IN SOME SPECIES THAN IN OTHERS?

Animal species differ in the variability of their clutch sizes, as well as in mean clutch sizes. This phenomenon is particularly obvious in lizards, where virtually invariant clutch sizes have evolved independently in at least 23 lineages in seven families. Reduced variance in clutch size may arise either as an adaptation (because females with less variable clutch sizes have higher fitness) or as an indirect by-product of selection on other life-history characteristics. Comparative data on Australian scincid lizards indicate that variance in clutch sizes is lowest among species with low mean clutch sizes, small body sizes and a low variance in body sizes of adult females. Phylogenetic analysis shows that evolutionary decreases in the variance of clutch size have accompanied decreases in mean clutch sizes and decreases in the variance of adult female body sizes. Tropical lizards may also exhibit lower variance in clutch size. Most of these characteristics are correlated in occurrence, and may be allometrically tied to small body size. Hence, low variance in clutch size may be a consequence of allometric effects on a correlated suite of life-history characteristics. Exceptions to the general patterns noted above—especially, lizard species with invariant clutch sizes but large body sizes—may be due to loss of genetic variance for clutch sizes in lineages that have passed through a “bottleneck” of small body sizes and hence, low variance in clutch sizes.     

Onset of maturity in male southern rock lobsters Jasus edwardsii in Tasmania,Australia

Summary

The reproductive contribution of male southern rock lobsters Jasus edwardsii has recently become of interest for managing this fishery as changes to the length of seasonal closures have shifted fishing effort towards males. To assess size at onset of maturity, a total of 387 male southern rock lobsters Jasus edwardsii were collected using baited traps from four of the eight fishery regions around the Tasmanian coast during July and August 2000. Sizes at onset of maturity in these samples were estimated by examining the vasa deferentia for evidence of the commencement of spermatid production. In addition, the total combined weight of the vasa deferentia and testis was compared with body size (as a gonad somatic index, GSI). Estimates of size at onset of maturity from the two methods were similar, although estimates from GSI appeared to have broader confidence limits. Although these physiological indices of maturity do not necessarily translate to functional maturity, they indicate that maturity is well below that of the legal minimum size of 110 mm carapace length used for managing the fishery (all estimates by either method for length at 50% sample maturity ≤70 mm carapace length). Production of sperm by males occurs below the size at onset of maturity of females from the same region and spatial patterns in maturity estimated by onset of spermatid production were similar to those of females.     

The problem of optimal clutch size in a tritrophic system: the oviposition strategy of the thistle gallfly Urophora cardui (Diptera,Tephritidae)

The problem of optimal clutch sizes is a central theme in life history theory. Optimal allocation of eggs is especially complicated for insects in tritrophic systems. In this study we analyze some of the processes determining clutch sizes of the thistle gallfly Urophora cardui, a monophagous tephritid fly associated with Cirsium arvense. U. cardui forms multilocular shoot galls, which vary broadly in their size and number of their gall cells. We investigate various fitness consequences of gall size. An analysis of the number of cells per gall (which is correlated with gall diameter and gall weight) showed that in U. cardui there is mutual facilitation rather than larval competition. Increasing numbers of larvae per gall led to a decreasing mortality and increasing larval weight. Larval weight in turn was positively correlated with the probability of survival to adulthood and with adult weight and fecundity. Thus, all fitness parameters measured favoured large galls. Clutch sizes in oviposition experiments were distinctly larger than the number of gall cells of field populations and in cage experiments, suggesting high mortality of eggs and/or early larval instars. There was a significant relationship between the internal structure (i.e., the size of the growing point) of the bud and clutch size, suggesting that U. cardui females are able to measure bud quality and adapt clutch sizes accordingly. Clutch size was positively correlated with the female's age at first oviposition and negatively with the number of previous ovipositions and previously laid eggs. Since the potential egg capacity per female is higher than the average number of larvae it is likely to produce during its short adult lifespan, U. cardui females tend to be time-limited rather than egglimited, which might favour large clutches once an appropriate oviposition site has been located. As the development of the gall and hence the fate of a clutch depends on a number of unpredictable factors, exclusive concentration of eggs in a few large clusters would involve risks which could be avoided by increasing the number of clutches. Therefore we interpret the high variation of clutch sizes in U. cardui as a mixed strategy of bet hedging and gambling.