Reproductive cycle of Golfingia margaritacea,a bipolar sipunculan,in subantarctic water

The reproductive cycle of the bipolar sipunculan Golfingia margaritacea collected from Ushuaia, Argentina (54°49 S; 68°13W) in 1986 and 1987 is reported. The population sampled showed a sex ratio of 1:1. Males, however, frequently outnumbered females. Measurements of coelomic oocytes delineated an annual reproductive cycle. Small oocytes were present in the coelom all year: a subpopulation began to increase size about the middle of spring and reached its maximum size in late summer. Spawns may occur from the end February to April. May is characterized by resorption of large oocytes. Monthy monitoring of the gonads revealed mitotic activity and a continuous release of gametocytes to the coelom. Parasites were detected in the gonads. Large coelomic oocytes were also present in the coelom during the same season in the Northern and Southern Hemispheres, but egg sizes were significantly larger in boreal populations.     

Gonadal development and infrequent sex change in a population of the humbug damselfish,Dascyllus aruanus,in continuous coral-cover habitat

The expression of protogyny often differs among populations of a single species. The humbug damselfish, Dascyllus aruanus (Linnaeus), typically lives in spatially discrete groups of several individuals around live coral colonies and has been reported to be protogynous. The size and sexual composition of groups of this species, however, vary with the size of the coral patch or the degree of coral cover. The present study examined gonadal development and sexual pattern in a population of D. aruanus living in continuous coral-cover habitat. Fish were collected in Tumon Bay, Guam (13°31′N; 144°47′E), during a 2-week period in July and August 1996. Reflecting their distributions, small fish (n=54) were collected at depths of 1–2 m over continuous Porites spp. patches, and large fish (n=56) were collected at depths of 2–3 m over a continuous Acropora spp. patch. Gonadal development was examined histologically. Gonads of D. aruanus first developed an ovarian lumen and primary-growth stage oocytes. From this ovarian state or from more developed ovaries with cortical-alveolus stage oocytes, some gonads developed into testes through degeneration of oocytes and development of spermatogenic tissue. Developing spermatogenic tissue only occurred in gonads having pre-cortical alveolus stage oocytes, and the size of individuals with gonads of this type overlapped mainly with the size range of individuals with gonads containing only pre-vitellogenic oocytes (i.e. immature females). This suggested that functional female-to-male sex change was rare or absent in this population of D. aruanus. Moreover, the different habitat distributions of small and large fish suggest that sex determination in immature fish occurs in the absence of adults.     

Reproduction and sex reversal of the solitary vermetid gastropod<Emphasis Type="Italic"> Serpulorbis arenarius</Emphasis>

The reproductive biology of the Mediterranean vermetid gastropod Serpulorbis arenarius, a subtidal sessile solitary species that lives in shady rocky communities, was studied at two localities south-east of the Iberian Peninsula. Its population density is usually low, with specimens sometimes sparsely distributed. Its shell often appears to be covered by epibiotic organisms. The sex ratio is biased toward males (63%) in the populations of SE Spain, and the results point towards hermaphroditism with alternating sexuality. The results obtained suggest that all specimens are first males, change to females under some as yet unknown condition and, after the ovogenesis cycle has been completed, revert to the original sexual state (male). The possible adaptive forces that favour hermaphroditism in this species are discussed, and we hypothesise that the seizure of a spermatophore by males may stimulate sex change to female. Brooding females (51% of females studied) were found from March to December. In males, histological study of testes revealed that spermatogenesis seems to occur throughout the year. Females brooded between 5 and 28 (=15) egg capsules, which were attached by stalks to the inner surface of the shell. It is noteworthy that the gonads of two of the specimens brooding egg capsules were developing testes. The newly laid egg capsules were filled with an organic intracapsular matrix, in which 40–153 (=94) eggs were embedded. The mean diameter of the uncleaved ova was 352.5 µm. The organic intracapsular matrix and some nurse eggs were used as an extra-embryonic food source. Hatching occurred at an advanced veliger stage, immediately followed by metamorphosis. Therefore, the larval development of S. arenarius is of a lecithotrophic intracapsular type, with a short, free-swimming/crawling phase.Communicated by O. Kinne, Oldendorf/Luhe     

Reproductive cycle of two populations of Ophionereis schayeri (Ophiuroidea) in New South Wales

Reproduction of two populations of Ophionereis schayeri (Müller and Troschel) in New South Wales was investigated from November 1991 through January 1993. The reproductive cycle was documented by histological examination of the gonads and by image analysis to determine the oocyte size-frequency distributions. Both populations of O. schayeri were mature in December and January and had a major summer spawning period between January and February. Thereafter, the condition of the gonads exhibited interindividual variability, with asynchronous low-intensity gamete release through August. O. schayeri has the potential to spawn for up to 8 mo of the year. This breeding pattern with synchronous spawning in summer and asynchronous gamete release through winter was similar in both populations. During autumn and winter, the gonads contained developing gametes and advanced gametes ready for spawning. The eggs spawned by O. schayeri during these seasons started their growth during the previous spring vitellogenic period, and continued to grow through summer. Upon reaching full size, they were stored for spawning outside the main breeding period. Spring is marked by increased spermatogenic and vitellogenic activity followed by maturation and spawning. Throughout their development, the oocytes of O. schayeri are surrounded by a follicle consisting of periodic acid-Schiff-positive (PAS+) haemal fluid and PAS+ yolk by the oocytes. In the testes, the haemal sinus projects into the centre of the spermatocyte columns, thus appearing to play a nutritive role in support of both oogenesis and spermatogenesis. Late vitellogenic oocytes of O. schayeri are firmly anchored to the germinal epithelium by an attachment complex consisting of specialised attachment cells and basophilic strands that radiate between the oolemma and the nucleus. Spawning is associated with rupture of the follicles, which remain as prominent, empty U-shaped profiles in the ovaries. O. schayeri produces copious numbers of 200 m-diam oocytes, suggesting that this species is a broadcast spawner and that it has a modified ophiopluteus or vitellaria larva.     

Reproduction and reproductive variability in the coral Pocillopora verrucosa from the Republic of Maldives

The sexuality, reproductive mode, and timing of reproduction of Pocillopora verrucosa from the Republic of Maldives, Indian Ocean, were assessed using serial histological sections. These showed that P. verrucosa is an annual simultaneous hermaphrodite, with gonads arranged in two opposing arcs of alternating testes and ovaries, six gonads in each arc. No planulae were observed in any dissection or histological analysis carried out, therefore making brooding unlikely. Broadcast spawning is inferred from the disappearance of mature gametes from samples collected between late March and April 1991. Mean oocyte size at spawning was 53.5 m and mean potential fecundity was 7300 ooctyes cm^-2yr^-1. The reproductive pattern of P. verrucosa in the Maldives is compared to that of the same species in different locations, the short breeding season in March to April occurring earlier in the Maldives than in Red Sea populations. The maximum mean oocyte diameter found in Maldivian specimens was much smaller than elsewhere. The year-to-year variation in numbers of oocytes/polyp within single colonies, and the variation between colonies within the population was significant (p<0.01). Therefore, it is possible that fecundity is not a good variable to use when monitoring stress on coral reefs unless larger numbers of estimates can be routinely made. Some colonies contained immature oocytes at spawning that were not released and continued to grow to approximately twice the size of the spawned oocytes that were presumed to be mature. These unspawned oocytes were oosorbing, and were characterised by the presence of zooxanthellae and large numbers of vacuoles in their cytoplasm. This gradual expansion and oosorption of unspawned oocytes has not been documented previously for corals.     

Reproductive cycle of the endemic cassiduloid<Emphasis Type="Italic"> Cassidulus mitis</Emphasis> (Echinoidea: Cassiduloida) on the Brazilian coast

Cassidulus mitis Krau, 1954 is an endemic species from Rio de Janeiro, Brazil. To study the reproductive cycle of this species, samples were collected at Praia Vermelha, in Guanabara Bay, from October 1998 to April 2000. The sex ratio for C. mitis was 1:1, and reproduction occurred throughout the year. The gonadal index of males was greater than that of females. Five gametogenic stages were identified in males: early growth, premature, mature, partially spawned and early growth with a partially spawned stage. There were no spent or recovery stages such as those seen in females. Females had six gametogenic stages: recovery, early growth, premature, mature, partially spawned and spent stage; there was no early growth with a partially spawned stage as there was in males. The mean oocyte diameter was 382 m (SD=49 m), and the mean number of juveniles per female was 100 individuals (SD=108). The anterior gonads were frequently smaller than the posterior ones, and there were differences in the gametogenic stages between the anterior and posterior gonads. First sexual maturation occurred in individuals with a diameter of 17 mm. The intestinal wet weight was two- to threefold higher than the test wet weight. This may indicate that C. mitis uses its intestine as a weight belt to avoid displacement from the substratum.Communicated by P.W. Sammarco, Chauvin     

Reproduction of<Emphasis Type="Italic"> Ophiactis resiliens</Emphasis> (Echinodermata: Ophiuroidea) in New South Wales with observations on recruitment

Reproduction of the ophiuroid Ophiactis resiliens Lyman 1879, a common species in the shallow waters around Australia, was examined over 1 year in two populations near Sydney, New South Wales. Gametogenesis was documented by histological examination of the gonads. Vitellogenesis in females and spermatocyte proliferation in males started in February and March. O. resiliens had an annual spawning period from May to September/October with peak gamete release from May to July. Thereafter, spawning occurred in some specimens through November. Gravid gonads were present for approximately 2 months longer in O. resiliens from Clovelly Bay than in those from Little Bay. At both sites spawning was episodic and gametogenesis continued through most of the spawning season. Most specimens (50–100%) collected from November to March had indeterminate/recovery stage gonads, the sex of which, could not be determined. The timing of gametogenic renewal during these months varied among individuals. O. resiliens has a 3-month planktonic stage and the presence of juveniles, 0.5 –1.5 mm disk diameter, in the coralline algal turf at Little Bay from February to May, suggests that settlement occurs in summer and autumn at this site. In contrast, juveniles were rarely encountered in coralline turf at Clovelly.     

Observations on the reproduction and postlarval morphology of pourtalesiid sea urchins in the Rockall Trough area (N.E. Atlantic Ocean)

Gametogenesis was studied histologically in the deep-sea species Pourtalesia jeffreysi (Wyville Thomson), P. miranda (A. Agassiz) and Echinosigra phiale (Wyville Thomson) collected during 1973–1983 from various positions at depths of 1 040 to 2 921 m in the Rockall Trough. The gonads are small and no synchrony in terms of the stage of gametogenic development was evident in females of the latter two species collected at different times of the year, indicating nonseasonal breeding. Males appear to be ripe at all times of the year. Follicle cells were well developed around the developing oocytes. Both the size range of the yolky ripe eggs (173 to 357 m diam) and potential fecundity within the range of 1 000 to 4 000 oocytes per individual, are indicative of an abbreviated, indirect, lecithotrophic mode of development occurring in all three species. Pourtalesiid postlarvae occurred also in fine-meshed trawl hauls, and their morphology is described. It was not possible to distinguish between P. miranda and E. phiale at sizes <3 mm in length.     

Bi-directional sex change in a coral-dwelling goby

Bi-directional sex change has recently been reported among obligate coral-dwelling gobies of the genus Gobiodon. However, neither the functional role of this pattern of sex change nor the frequency of sex change in either direction in natural populations is known. We investigated the social structure and pattern of sex change of Gobiodon histrio at Lizard Island on the Great Barrier Reef. The social structure of G. histrio within coral colonies usually consisted of a single juvenile or a heterosexual adult pair. The size of adult social groups was not constrained by coral colony size. In contrast to expectations for pair-forming species, G.␣histrio was primarily a protogynous hermaphrodite. All immature G. histrio were females and sex change from female to male occurred readily when two mature females were placed in a coral colony. In addition, male G. histrio were able to change back to females when two mature males were placed in a coral. Sex change from female to male, however, occurred with over twice the frequency of sex change from male to female. Where two males were placed in a coral colony, heterosexual pairs were most frequently re-established by immigration of females from outside the treatment population. This pattern might be predicted if sex change from male to female is more expensive than sex change from female to male for G. histrio. Where sex change is expensive, movement may be favoured over sex change, particularly where coral densities are high and movement among corals incurs little mortality risk. Received: 10 November 1997 / Accepted after revision: 16 May 1998     

Long-term changes in reproductive patterns of the holothurian<Emphasis Type="Italic"> Oneirophanta mutabilis</Emphasis> from the Porcupine Abyssal Plain

Time-series studies (1989–2002) in the NE Atlantic have shown large-scale changes in the composition and structure of the benthic community on the Porcupine Abyssal Plain. Radical changes in the abundance of some species in 1996 led to a significant shift in the way in which organic matter was reprocessed at the seabed. This article examines the reproductive processes of the holothurian Oneirophanta mutabilis collected during the time series. The reproductive biology of O. mutabilis is reviewed. No males were evident in any of the samples. The sex ratio (females: no gender) was biased significantly towards no-gender individuals. The maximum egg size was 650 m and there was no evidence of synchrony in reproduction. Significant changes in the oocyte-size distribution and the fecundity of O. mutabilis were noted through time, coinciding with the time of greatest faunal change in the benthic community. There was an increase in the proportion of previtellogenic oocytes and a decrease in the proportion of mature vitellogenic oocytes in 1997 and 1998, in parallel with a significant decrease in fecundity of the post-1996 samples. Samples from 2002 showed a reversal in the reproductive trends, with an increase in the proportion of mature vitellogenic oocytes and fecundity. The results are discussed in relation to large changes in abundance of the epibenthos on the Porcupine Abyssal Plain. It is suggested that the superabundance of certain megafaunal species on the abyssal seafloor affected the availability of trophic resources for O. mutabilis, leading to the changes in its reproductive effort.     

Gene flow among giant clam (Tridacna gigas) populations in Pacific does not parallel ocean circulation

Tridacna gigas (L.) populations were sampled from 13 locations throughout the west Pacific in 1990–1991 and screened for allozyme variation at seven polymorphic loci. Significant genetic differences were found between the few large stocks of T. gigas remaining in the world. Cluster analysis identified two major regional groups of populations: a Central Pacific group consisting of Kiribati and the Marshall Islands, and a West Pacific group comprising all other populations. Within the latter group, the Great Barrier Reef populations were significantly differentiated from those in the Solomon Islands. The main routes of gene flow did not run parallel with known major surface currents as might be expected, but crossed them. Gene exchange was limited east-west between the Central and West Pacific groups and the GBR and the Solomon Islands. The lack of correspondence between the major surface currents and the routes of gene flow suggests that the genetic structure of T. gigas reflects historical patterns of migration that no longer occur, rather than presentday dispersal. These findings raise fundamental questions concerning the origin and maintenance of genetic diversity in Pacific faunas, and have important implications for the mariculture and conservation of T. gigas.     

Reproductive patterns of scleractinian corals in the northern Red Sea

The majority of published accounts on scleractinian coral reproduction are from the tropical Pacific and Caribbean, with very little information known about Red Sea species. This report examines variation in reproductive mode in 24 species of hermatypic corals (belonging to seven families) in the Gulf of Eilat, Red Sea. Eighteen species are hermaphroditic broadcasters, two are hermaphroditic brooders and three are gonochoric broadcasters. In the Pocilloporidae, the gonads project into the body cavity, while in the other six families the gonads reside inside the mesenteries. The number of gonads per polyp in broadcasting species follows family or genus lines. Fecundity (eggs per polyp) increases with polyp size. Brooding species usually exhibit one or two gonads per polyp and each gonad contains only one to three oocytes. Oocyte size varies widely and does not relate to mode of reproduction. The largest oocytes (diameter = 450 μm) occur in the brooding coral Alveopora daedalea and in broadcasting species of the genus Acropora (diameter = 420 μm). Gonad morphology and gonochorism versus hermaphroditism appear to be constrained phylogenetically at the family or genus level. Lastly, this report compares the data presented for Red Sea scleractinian species with the data available on scleractinian corals from other geographical regions. Received: 2 February 1993 / Accepted: 9 March 1998     

Sexual maturity and hermaphroditism of the red porgy Pagrus pagrus (Teleostei: Sparidae)

Gonadal changes related to sexual maturity were examined by histology in reared populations of the red porgy Pagrus pagrus L., aged 0.5 to 6.5 years. More than 50% of red porgies were mature at the age of 4 years although some individuals were already mature at the age of 3, as indicated by changes in the gonadosomatic index, plasma levels of vitellogenin, 17β-estradiol and testosterone in females and males. Sex frequency distribution analysis in relation to the age and histological analysis of the gonads indicated the presence of males originated from the sex change of adult functional females. P. pagrus is a protogynous species. Some females had not changed sex up to the age of 6.5 years. Received: 16 July 1998 / Accepted: 29 April 1999     

Geographic and reproductive isolation in the marine Harpacticoid Copepod Tisbe

Within the genus Tisbe (Copepoda: Harpacticoida), there is a wide range of reproductive isolation. In the European area, in addition to species where (between geographic populations) the reproductive barrier is practically absent, there are others which show a varying degree of incompatibility. The present paper deals with the results of cross-breeding experiments carried out between American and European populations. The respective intrapopulation crosses served as controls. Transatlantic crosses have been attempted with the following species: T. lagunaris, T. bulbisetosa, T. holothuriae, T. battagliai, and T. clodiensis. The American and European populations of the first 4 species are interfertile, and the F₁ hybrids produce viable offspring. Moreover, the interpopulation crosses, compared to the controls, exhibit a slight superiority whose nature is, however, still unclear. The fifth species, T. clodiensis, behaves differently. The absence of viable offspring from interpopulation crosses, or the death of the hybrids before reaching the adult stage, indicate that, in this species, the barrier to gene flow is complete. Populations of T. clodiensis show, in addition, signs of relative intraspecific incompatibility. The problem of the different degree of reproductive isolation in the genus Tisbe, with special regard to the role played by ecological factors in determining the isolation patterns, is discussed.     

Plasma levels of androgens and gonadal development of breeders and helpers in the bell miner (Manorina melanophrys)

Plasma concentration of androgens and gonad development were studied in wild cooperatively breeding bell miners (Manorina melanophrys) of different age, sex and social status. Plasma levels of androgens increased with age of birds. For sexually mature (SM) bell miners, androgen levels were higher in breeders than in nonbreeding (NB) helpers (P < 0.005), and they were marginally higher in females than in males (P < 0.10). Thus female breeders showed the highest plasma levels of androgens. This pattern may be explained by an asymmetry between the sexes in the levels of competition needed to reach and defend breeding status; the degree of intra-sexual competition seems to be higher among female bell miners than among males. In general, male breeders had larger gonads than male SMNB helpers; male breeder gonads produced spermatozoa whereas most helpers' gonads did not. However, old (i.e. 34 months) SMNB male helpers had large gonads producing spermatozoa in spite of having low plasma levels of androgens. We suggest that young male bell miners may be under a voluntary mechanism of reproductive suppression perhaps favoured by inbreeding avoidance; although some aggressive interactions observed between very young helpers (about 2 months) and adult members of the breeding unit suggests that a certain degree of imposition also occurs at young ages. Older male helpers may avoid aggression with the male breeder and imposed reproductive suppression, hence the low circulating levels of androgens, but they have the potential to sneak extra-pair copulations. Gonad size decreased during the non-breeding season in both sexes.     

Divide or broadcast: Interrelation of asexual and sexual reproduction in a population of the fissiparous hermaphroditic seastar Nepanthia belcheri (Asteroidea: Asterinidae)

Asexual and sexual reproduction were studied in an intertidal population of Nepanthia belcheri (Perrier) at Townsville, Queensland, Australia, by regular sampling over a year (March 1976-March 1977) and by histological analysis of gonads. Fission reached a peak in early winter (April–June), when about 45% of the population showed evidence of recent fission. Propensity for fission was unrelated to longest arm length. Seven-armed seastars predominated in the population and these underwent fission in two stages to produce one 3-armed fragment and two 2-armed fragments. Fission planes were not related to numbers or positions of madreporites. Hermaphroditism was a normal sexual condition in the population. Almost all gonads contained oocytes, but some gonads functioned as ovaries (without spermatogenic tissue) while others functioned as testes. Seastars with mature ovaries were significantly larger than those with mature testes, indicating protandry, as in other hermaphroditic asteroids. However, fission complicates the pattern of gonad development by causing regression or retardation of gonads and by apparently having a masculinizing effect, so that ovaries may change to testes in fission products. There was a period of sexual reproduction in early summer (October–November). This followed the period of intense fission and regeneration, and a population change from predominately functional females to males. Consequently there was an extreme imbalance against mature females at sexual reproduction, further reducing potential fecundity. Thus, sexual reproduction was very subordinate to fission as the means of recruitment. The 450 m diam eggs probably give rise to pelagic lecithotrophic development and, if this is the case, N. belcheri retains the advantage of complementing reliable recruitment from fission with a dispersive phase. The combination of fission and hermaphroditism is particularly advantageous for a very sparse dispersal of larvae, as a functionally dioecious population may develop from one larva settling in a new locality.     

Genetic divergence between east and west Atlantic populations of<Emphasis Type="Italic"> Actinia</Emphasis> spp. sea anemones (Cnidaria: Actiniidae)

The sea anemone Actinia equina was considered a highly variable species with a wide geographical distribution, but molecular systematic studies have shown that this wide distribution may be the result of the lumping of cryptic species. In this work enzyme electrophoresis was used to analyse the genetic variability of A. equina from the Atlantic coasts of Europe and Africa, as well as the relationships between those populations and other species of the genus. Samples of A. equina from the United Kingdom and France were compared with supposedly conspecific populations from South Africa and a recently described species from Madeira, Actinia nigropunctata. The South African and Madeiran populations were genetically very divergent from each other (genetic identity, I=0.15), as well as from the A. equina population from the United Kingdom and from the other species studied (I between 0.23 and 0.78), indicating that the African population of A. equina belongs to a different species. High genetic similarities were found between west Atlantic A. bermudensis and east Atlantic A. sali, in spite of the great geographical distance between them, and between the European A. equina and A. prasina. The results confirm the presence of the new species A. nigropunctata from Madeira and suggest a new species within the A. equina species group, from South Africa.Communicated by O. Kinne, Oldendorf/Luhe     

Earlier sex change in infected individuals of the protogynous reef fish<Emphasis Type="Italic"> Thalassoma bifasciatum</Emphasis>

Sex allocation theory for sequential hermaphrodites predicts the size at which an individual should change sex, given the different relationships between individual size and reproductive success in the two sexes. We studied a host-parasite system where the myxozoan Kudoa ovivora infects the ovaries of the reef fish Thalassoma bifasciatum, a protogynous sequential hermaphrodite. The parasite sporulates in the hosts eggs and renders them infertile. It is thus expected to reduce the females reproductive success, and could thereby influence host sex change. We present data from marked fish we observed in the field over 4 months. The data suggest that females infected with Kudoa ovivora have a lower reproductive success, change sex earlier and at a smaller size than uninfected females. These results are in agreement with predictions from sex allocation theory, and provide the first example of a possible parasitic influence on the sex allocation of its host.Communicated by P. Heeb     

Reproductive strategies of two deep-sea gastropod species from the Porcupine Seabight (Northeast Atlantic)

Two species of small gastropods (<6 mm in length), Amphissa acutecostata (Philippi, 1844) and Gymnobela subaraneosa (Dautzenberg and Fischer, 1896), widely distributed in the northeast Atlantic, were found in large numbers in the Porcupine Seabight (Northeast Atlantic). Except for some aspects of taxonomy and distribution, as well as some data on larval development, the biology of these species is unknown. This study describes basic aspects of the life-history strategies of both species. Histological studies showed that oocyte and sperm development in both species was similar to the gametogenetic patterns observed in other deep-sea gastropods. In females, oogonia proliferated in the germinal epithelium and developed into previtellogenic oocytes (30–40 m), which grew into vitellogenic primary oocytes. Vitellogenic oocytes were covered by a thin layer of follicle cells involved in the vitellogenic processes. The maximum size for mature oocytes was 99.06 m for A. acutecostata and 114.82 m for G. subaraneosa. In A. acutecostata most of the volume of the ovary was occupied by previtellogenic and early vitellogenic oocytes, whereas in G. subaraneosa most of the volume was filled by large vitellogenic oocytes. Both species showed quasi-continuous production of oocytes. The oocyte size-frequency diagrams suggested a continuous release of a small number of oocytes throughout the year for A. acutecostata, and asynchronous periodic spawning events for G. subaraneosa. Gonad development and gametogenesis could be strongly affected by presence of parasites in one of the populations of G. subaraneosa.Communicated by J.P. Thorpe, Port Erin     

Ecological investigations of the zooplankton community of balsfjorden,northern norway: The genital system in Calanus finmarchicus and the role of gonad development in overwintering strategy

The structure of the reproductive system in Copepodite Stages IV and V of the marine copepod Calanus finmarchicus (Gunnerus) is described, together with seasonal variations in gonad development and sex ratio, from Balsfjorden (69°21N; 19°06E), a subarctic fjord in northern Norway. The genital system in Copepodite Stage IV consists of a small gonad with two genital ducts. The gonad of Copepodite Stage V in its most immature condition resembles that seen in Copepodite Stage IV, whereas the sex of Stage V copepodites with mature gonads can be easily and reliably determined. The genital ducts are the key characters for separating males and females in Copepodite Stage V; potential males have one and potential females two genital ducts connecting the gonad to the genital opening on the first urosome segment. Adults males were rare compared to females, and were only in the majority at the end of January and beginning of February. The sex-ratio also favoured females in Copepodite Stage V. Seasonal peaks in the proportion of adult males reflected peaks of males in Copepodite Stage V, suggesting that seasonal variations in the adult sex-ratio are a direct result of a situation which has already been determined in Copepodite Stage V. The seasonal variation in gonad development in Copepodite Stages IV and V of C. finmarchicus reveals that the size of maturity of the gonad varies throughout the year. This study indicates that visible sex-differentiation in Stage V and moulting into adults occurs in Balsfjorden at least 2 months before spawning in April. Phytoplankton levels are immeasurable before the spring diatom increase starts at the end of March, implying that development and maturation of gonads are dependent upon internal energy resources, whereas the final act of spawning seems to require energy input from phytoplankton.