Experimental research on the development of Ceratium hirundinella O.F.Muller [Translation from: Z.Bot. 14, 337-371, 1922]

The most important aim of this study lay in filling in the great gap in our knowledge of the processes of germination in the Ceratium cyst and the early developmental stages in the standing stock of Ceratium hirundinella. contained rich cysts, we now succeeded extraordinarily well in pursuing the consistent development of Ceratium from the cyst to the completed cell. A series of experiments were carried out on the cysts and the juvenile stages of Ceratium, which showed very interesting results. The author presents in a general descriptive part the normal processes of germination in Ceratium cysts and the development of the juvenile stages in order to show in an experimental part the changes in form of C. hirundinella under the influence of temperature, light and varying salinities.

Estimates of body sizes at maturation and at sex change, and the spawning seasonality and sex ratio of the endemic Hawaiian grouper (Hyporthodus quernus, F. Epinephelidae)

A case study of the reproductive biology of the endemic Hawaiian grouper or hapu’upu’u (Hyporthodus quernus) is presented as a model for comprehensive future studies of economically important epinephelid groupers. Specimens were collected throughout multiple years (1978–81, 1992–93, and 2005–08) from most reefs and banks of the Northwestern Hawaiian Islands. The absence of small males, presence of atretic oocytes and brown bodies in testes of mature males, and both developed ovarian and testicular tissues in the gonads of five transitional fish provided evidence of protogynous hermaphroditism. No small mature males were collected, indicating that Hawaiian grouper are monandrous (all males are sex-changed females). Complementary microscopic criteria also were used to assign reproductive stage and estimate median body sizes (L50) at female sexual maturity and at adult sex change from female to male. The L50 at maturation and at sex change was 580 ±8 (95% confidence interval [CI]) mm total length (TL) and 895 ±20 mm TL, respectively. The adult sex ratio was strongly female biased (6:1). Spawning seasonality was described by using gonadosomatic indices. Females began ripening in the fall and remained ripe through April. A February–June main spawning period that followed peak ripening was deduced from the proportion of females whose ovaries contained hydrated oocytes, postovulatory follicles, or both. Testes weights were not affected by season; average testes weight was only about 0.2% of body weight—an order of magnitude smaller than that for ovaries that peaked at 1–3% of body weight. The species’ reproductive life history is discussed in relation to its management.

Advances in methods for determining fecundity: application of the new methods to some marine fishes

Estimation of individual egg production (realized fecundity) is a key step either to understand the stock and recruit relationship or to carry out fisheries-independent assessment of spawning stock biomass using egg production methods. Many fish are highly fecund and their ovaries may weigh over a kilogram; therefore the work time can be consuming and require large quantities of toxic fixative. Recently it has been shown for Atlantic cod (Gadus morhua) that image analysis can automate fecundity determination using a power equation that links follicles per gram ovary to the mean vitellogenic follicular diameter (the autodiametric method). In this article we demonstrate the precision of the autodiametric method applied to a range of species with different spawning strategies during maturation and spawning. A new method using a solid displacement pipette to remove quantitative fecundity samples (25, 50, 100, and 200 milligram [mg]) is evaluated, as are the underlying assumptions to effectively fix and subsample the ovary. Finally, we demonstrate the interpretation of dispersed formaldehyde-fixed ovarian samples (whole mounts) to assess the presence of atretic and postovulatory follicles to replace labor intensive histology. These results can be used to estimate down regulation (production of atretic follicles) of fecundity during maturation.

Sexual differentiation and gonad development in striped mullet (Mugil cephalus L.) from South Carolina estuaries*

This study examined the sexual differentiation and reproductive dynamics of striped mullet (Mugil cephalus L.) in the estuaries of South Carolina. A total of 16,464 specimens were captured during the study and histological examination of sex and maturity was performed on a subsample of 3670 fish. Striped mullet were sexually undifferentiated for the first 12 months, began differentiation at 13 months, and were 90% fully differentiated by 15 to 19 months of age and 225 mm total length (TL). The defining morphological characteristics for differentiating males was the elongation of the protogonial germ tissue in a corradiating pattern towards the center of the lobe, the development of primary and secondary ducts, and the lack of any recognizable ovarian wall structure. The defining female characteristics were the formation of protogonial germ tissue into spherical germ cell nests, separation of a tissue layer from the outer epithelial layer of the lobe-forming ovarian walls, a tissue bud growing from the suspensory tissue that helped form the ovary wall, and the proliferation of oogonia and oocytes. Sexual maturation in male striped mullet first occurred at 1 year and 248 mm TL and 100% maturity occurred at age 2 and 300 mm TL. Female striped mullet first matured at 2 years and 291 mm total length and 100% maturity occurred at 400 mm TL and age 4. Because of the open ocean spawning behavior of striped mullet, all stages of maturity were observed in males and females except for functionally mature females with hydrated oocytes. The spawning season for striped mullet recruiting to South Carolina estuaries lasts from October to April; the majority of spawning activity, however, occurs from November to January. Ovarian atresia was observed to have four distinct phases. This study presents morpholog ical analysis of reproductive ontogeny in relation to size and age in South Carolina striped mullet. Because of the length of the undifferentiated gonad stage in juvenile striped mullet, previous studies have proposed the possibility of protandric hermaphrodism in this species. The results of our study indicate that striped mullet are gonochoristic but capable of exhibiting nonfunctional hermaphroditic characteristics in differentiated mature gonads.

Reproductive biology of female Rikuzen sole (Dexistes rikuzenius)*

The annual ovarian cycle, mode of maturation, age at maturity, and potential fecundity of female Rikuzen sole (Dexistes rikuzenius) from the North Pacific Ocean off the coast of Japan were studied by 1) histological examination of the gonads, 2) measurement and observation of the oocytes, and 3) by otolith aging. The results indicated that ovulation occurs from September to December and peaks between September and October. Vitellogenesis began again soon after the end of the current season. Maturity was divided into eight phases on the basis of oocyte developmental stages. Mature ovaries contained developing oocytes and postovulatory follicles but no recruiting oocytes, indicating that this species has group-synchronous ovaries and is a multiple spawner. Almost all females matured first at an age of 1+ year and spawned every year until at least age 8+ years. Potential fecundity increased exponentially with body length and the most fecund fish had 15 times as many oocytes as the least fecund fish. Potential fecundity and relative fecundity were both positively correlated with age from 1 to 6+ years, but were negatively correlated, probably because of senescence, in fish over 7 years. These results emphasize that the total productivity of a D. rikuzenius population depends not only on the biomass of females older than 1+ but also on the age structure of the population.

Movements and spawning of white marlin (Tetrapturus albidus) and blue marlin (Makaira nigricans) off Punta Cana, Dominican Republic

With a focus on white marlin (Tetrapturus albidus), a concurrent electronic tagging and larval sampling effort was conducted in the vicinity of Mona Passage (off southeast Hispaniola), Dominican Republic, during April and May 2003. Objectives were 1) to characterize the horizontal and vertical movement of adults captured from the area by using pop-up satellite archival tags (PSATs); and 2) by means of larval sampling, to investigate whether fish were reproducing. Trolling from a sportfishing vessel yielded eight adult white marlin and one blue marlin (Makaira nigricans); PSAT tags were deployed on all but one of these individuals. The exception was a female white marlin that was unsuitable for tagging because of injury; the reproductive state of its ovaries was examined histologically. Seven of the PSATs reported data summaries for water depth, temperature, and light levels measured every minute for periods ranging from 28 to 40 days. Displacement of marlin from the location of release to the point of tag pop-up ranged from 3l.6 to 267.7 nautical miles (nmi) and a mean displacement was 3.4 nmi per day for white marlin. White and blue marlin mean daily displacements appeared constrained compared to the results of other marlin PSAT tagging studies. White marlin ovarian sections contained postovulatory follicles and final maturation-stage oocytes, which indicated recent and imminent spawning. Neuston tows (n=23) yielded 18 istiophorid larvae: eight were white marlin, four were blue marlin, and six could not be identified to species. We speculate that the constrained movement patterns of adults may be linked to reproductive activity for both marlin species, and, if true, these movement patterns may have several implications for management. Protection of the potentially important white marlin spawning ground near Mona Passage seems warranted, at least until further studies can be conducted on the temporal and spatial extent of reproduction and associated adult movement.

Reproductive biology of carpenter seabream (Argyrozona argyrozona) (Pisces: Sparidae) in a marine protected area

The carpenter seabream (Argyrozona argyrozona) is an endemic South African sparid that comprises an important part of the handline fishery. A three-year study (1998−2000) into its reproductive biology within the Tsitsikamma National Park revealed that these fishes are serial spawning late gonochorists. The size at 50% maturity (L50) was estimated at 292 and 297 mm FL for both females and males, respectively. A likelihood ratio test revealed that there was no significant difference between male and female L50 (P>0.5). Both monthly gonadosomatic indices and macroscopically determined ovarian stages strongly indicate that A. argyrozona within the Tsitsikamma National Park spawn in the astral summer between November and April. The presence of postovulatory follicles (POFs) confirmed a six-month spawning season, and monthly proportions of early (0−6 hour old) POFs showed that spawning frequency was highest (once every 1−2 days) from December to March. Although spawning season was more highly correlated to photoperiod (r = 0.859) than temperature (r = −0.161), the daily proportion of spawning fish was strongly correlated (r= 0.93) to ambient temperature over the range 9−22oC. These results indicate that short-term upwelling events, a strong feature in the Tsitsikamma National Park during summer, may negatively affect carpenter fecundity. Both spawning frequency and duration (i.e., length of spawning season) increased with fish length. As a result of the allometric relationship between annual fecundity and fish mass a 3-kg fish was calculated to produce fivefold more eggs per kilogram of body weight than a fish of 1 kg. In addition to producing more eggs per unit of weight each year, larger fish also produce significantly larger eggs.

Fecundity of shortspine thornyhead (Sebastolobus alascanus) and longspine thornyhead (S. altivelis) (Scorpaenidae) from the northeastern Pacific Ocean, determined by stereological and gravimetric techniques*

Fecundity was estimated for shortspine thornyhead (Sebastolobus alascanus) and longspine thornyhead (S. altivelis) from the northeastern Pacific Ocean. Fecundity was not significantly different between shortspine thornyhead off Alaska and the West Coast of the United States and is described by 0.0544 × FL3.978, where FL =fish fork leng th (cm). Fecundity was estimated for longspine thornyhead off the West Coast of the United States and is described by 0.8890 × FL3.249. Contrary to expectations for batch spawners, fecundity estimates for each species were not lower for fish collected during the spawning season compared to those collected prior to the spawning season. Stereological and gravimetric fecundity estimation techniques for shortspine thornyhead provided similar results. The stereological method enabled the estimation of fecundity for samples collected earlier in ovarian development; however it could not be used for fecundity estimation in larger fish.

On the trail of interannual variability in phytoplankton and climate [abstract and figures]

EXTRACT (SEE PDF FOR FULL ABSTRACT): A selective but nontheless real record of phytoplankton activity over the Santa Barbara Basin can be obtained from the underlying varved sediments. The phytoplankton groups preserved are: diatoms (frustrules and spores), silicoflagellates, dinoflagellates (cysts) and coccoliths.

Staging ovaries of Haddock (Melanogrammus aeglefinus): implications for maturity indices and field sampling practices

We build on recent efforts to standardize maturation staging methods through the development of a field-proof macroscopic ovarian maturity index for Haddock (Melanogrammus aeglefinus) for studies on diel spawning periodicity. A comparison of field and histological observations helped us to improve the field index and methods, and provided useful insight into the reproductive biology of Haddock and other boreal determinate fecundity species. We found reasonable agreement between field and histological methods, except for the regressing and regenerating stages (however, differentiation of these 2 stages is the least important distinction for determination of maturity or reproductive dynamics). The staging of developing ovaries was problematic for both methods partly because of asynchronous oocyte hydration during the early stage of oocyte maturation. Although staging on the basis of histology in a laboratory is generally more accurate than macroscopic staging methods in the field, we found that field observations can uncover errors in laboratory staging that result from bias in sampling unrepresentative portions of ovaries. For 2 specimens, immature ovaries observed during histological examination were incorrectly assigned as regenerating during macroscopic staging. This type of error can lead to miscalculation of length at maturity and of spawning stock biomass, metrics that are used to characterize the state of a fish population. The revised field index includes 3 new macroscopic stages that represent final oocyte maturation in a batch of oocytes and were found to be reliable for staging spawning readiness in the field. The index was found to be suitable for studies of diel spawning periodicity and conforms to recent standardization guidelines.

Reproductive potential of Yellowfin Tuna (Thunnus albacares) in the western Indian Ocean

The reproductive biology of Yellowfin Tuna (Thunnus albacares) in the western Indian Ocean was investigated from samples collected in 2009 and 2010. In our study, 1012 female Yellowfin Tuna were sampled: 320 fish on board a purse seiner and 692 fish at a Seychelles cannery. We assessed the main biological parameters that describe reproductive potential: maturity, spawning seasonality, fish condition, and fecundity. The length at which 50% of the female Yellowfin Tuna population matures (L50) was estimated at 75 cm in fork length (FL) when the maturity threshold was established at the cortical alveolar stage of oocyte development. To enable comparison with previous studies, L50 also was estimated with maturity set at the vitellogenic stage of oocyte development; this assessment resulted in a higher value of L50 at 102 cm FL. The main spawning season, during which asynchrony in reproductive timing among sizes was observed, was November–February and a second peak occurred in June. Smaller females (<100 cm FL) had shorter spawning periods (December to February) than those (November to February and June) of large individuals, and signs of skip-spawning periods were observed among small females. The Yellowfin Tuna followed a “capital-income” breeder strategy during ovarian development, by mobilizing accumulated energy while using incoming energy from feeding. The mean batch fecundity for females 79–147 cm FL was estimated at 3.1 million oocytes, and the mean relative batch fecundity was 74.4 oocytes per gram of gonad-free weight. Our results, obtained with techniques defined more precisely than techniques used in previous studies in this region, provide an improved understanding of the reproductive cycle of Yellowfin Tuna in the western Indian Ocean.

An atlas of reproductive development in rockfishes, genus Sebastes

The genus Sebastes consists of over 100 fish species, all of which are viviparous and long-lived. Previous studies have presented schemes on the reproductive biology of a single targeted species of the genus Sebastes, but all appear to possess a similar reproductive biology as evidenced by this and other studies. This atlas stages major events during spermatogenesis, oogenesis, and embryogenesis, including atresia, in six species of Sebastes (S. alutus, S. elongatus, S. helvomaculatus, S. polyspinis, S. proriger, and S. zacentrus). Our study suggests that the male reproductive cycle of Sebastes is characterized by 11 phases of testicular development, with 10 stages of sperm development and 1 stage of spermatozoa atresia. Ovarian development was divided into 12 phases, with 10 stages of oocyte development, 1 stage of embryonic development, and 1 stage of oocyte atresia. Embryonic development up to parturition was divided into 33 stages following the research of Yamada and Kusakari (1991). Reproductive development of all six species examined followed the developmental classifications listed above which may apply to all species of Sebastes regardless of the number of broods produced annually. Multiple brooders vary in that not all ova are fertilized and progress to embryos; a proportion of ova are arrested at the pre-vitellogenic stage. Reproductive stage examples shown in this atlas use S. elongates for spermatic development, S. proriger for oocyte development, and S. alutus for embryological development, because opportunistic sampling only permitted complete analysis of each respective developmental phase for those species. The results of this study and the proposed reproductive phases complement the recommended scheme submitted by Brown-Peterson et al. (2011), who call for a standardization of terminology for describing reproductive development of fishes.

Oogenesis and spawn formation in the invasive lionfish, Pterois miles and Pterois volitans

The Indo-Pacific lionfish, Pterois miles and P. volitans, have recently invaded the U.S. east coast and the Caribbean and pose a significant threat to native reef fish communities. Few studies have documented reproduction in pteroines from the Indo-Pacific. This study provides a description of oogenesis and spawn formation in P. miles and P. volitans collected from offshore waters of North Carolina, U.S.A and the Bahamas. Using histological and laboratory observations, we found no differences in reproductive biology between P. miles and P. volitans. These lionfish spawn buoyant eggs that are encased in a hollow mass of mucus produced by specialized secretory cells of the ovarian wall complex. Oocytes develop on highly vascularized peduncles with all oocyte stages present in the ovary of spawning females and the most mature oocytes placed terminally, near the ovarian lumen. Given these ovarian characteristics, these lionfish are asynchronous, indeterminate batch spawners and are thus capable of sustained reproduction throughout the year when conditions are suitable. This mode of reproduction could have contributed to the recent and rapid establishment of these lionfish in the northwestern Atlantic and Caribbean.

Sex-specific growth and mortality, spawning season, and female maturation of the stripey bass (Lutjanus carponotatus) on the Great Barrrier Reef

Sex-specific demography and reproductive biology of stripey bass (Lutjanus carponotatus) (also known as Spanish flag snapper, FAO) were examined at the Palm and Lizard island groups, Great Barrier Reef (GBR).Total mortality rates were similar between the sexes. Males had larger L∞ at both island groups and Lizard Island group fish had larger overall L∞. Female:male sex ratios were 1.3 and 1.1 at the Palm and Lizard island groups, respectively. The former is statistically different from 1, but is unlikely significantly different in a biological sense. Females matured on average at 2 years of age and 190 mm fork length at both locations. Female gonadal lipid body indices peaked from August through October, preceding peak gonadosomatic indices in October, November, and December that were twice as great as in any other month. However, ovarian staging revealed 50% or more ovaries were ripe from September through February, suggesting a more protracted spawning season and highlighting the different interpretations that can arise between gonad weight and gonad staging methods. Gonadosomatic index increases slightly with body size and larger fish have a longer average spawning season, which suggests that larger fish produce greater relative reproductive output. Lizard Island group females had ovaries nearly twice as large as Palm Island group females at a given body size. However, it is unclear whether this reflects spatial differences akin to those observed in growth or effects of sampling Lizard Island group fish closer to their date of spawning. These results support an existing 250 mm minimum size limit for L. carponotatus on the GBR, as well as the timing of a proposed October through December spawning closure for the fishery. The results also caution against assessing reef-fish stocks without reference to sex-, size-, and location-specific biological traits.

Testicular development in migrant and spawning bluefin tuna (Thunnus thynnus (L.)) from the eastern Atlantic and Mediterranean

Testis histological structure was studied in bluefin tuna (Thunnus thynnus) from the eastern Atlantic and Mediterranean during the reproductive season (from late April to early June). Testicular maturation was investigated by comparing samples from bluefin tuna caught on their eastward reproductive migration off Barbate (Strait of Gibraltar area) with samples of bluefin tuna fished in spawning grounds around the Balearic Islands. Histological evaluations of cross sections showed that the testis consists of two structurally different regions, an outer proliferative region where germ cells develop synchronously in cysts, and a central region made up of a well-developed system of ducts that convey the spermatozoa produced in the proliferative region to the main sperm duct. Ultrastructural features of the different stages of the male germ cell line are very similar to those described in other teleost species. The bluefin tuna testis is of the unrestricted spermatogonial testicular type, where primary spermatogonia are present all along the germinative portion of the lobules. All stages of spermatogenesis were present in the gonad tissue of migrant and spawning bluefin tuna, although spermatids were more abundant in spawning fish. The testis size was found to increase by a factor of four (on average) during migration to the Mediterranean spawning grounds, whereas the fat bodies (mesenteric lipid stores associated with the gonads) became reduced to half their weight, and the liver mass did not change significantly with sexual maturation. Linear regression analysis of the pooled data of migrant and spawning bluefin tuna revealed a significant negative correlation between the gonad index (IG) and the fat tissue index (IF), and a weaker positive correlation between the gonad index (IG) and the liver index (IL). Our analyses indicate that the liver does not play a significant role in the storage of lipids and that mesenteric lipid reserves constitute an important energy source for gametogenesis in bluefin tuna.