Population dynamics of the commensal spider crab Inachus phalangium (Decapoda: Maiidae)

The ecology of the spider crab Inachus phalangium (Fabricius, 1775) (Decapoda: Maiidae) was studied in the field. I. phalangium inhabits the sublittoral on the sea anemone Anemonia sulcata Pennant. From July 1981 to April 1984 in the Mediterranean (Banyuls sur Mer, southern France) more than 3000 anemones were examined and ca 1800 I. phalangium were found on them. The population dynamics' generation cycles, reproductive activities and the dynamics of the sex ratio were investigated. The density of juveniles (crabs before the pubertal moult) on anemones changed in a yearly cycle from low in the first six months to very high in the second six months. The first occasional young crabs of a generation appeared in March/April (3rd and 4th decapodite stages) on the anemones. Their density increased enormously in the following months. The generation grew gradually on the anemones and moulted into puberty in September-January. Density of adults (crabs after the pubertal moult) on anemones changed in a yearly cycle from low to high from the summer to winter months. A new adult generation was recruited every autumn through the pubertal moult and disappeared in the following summer. Female reproductive activity continued throughout the year. Females carried several broods in succession, but the frequency of breeding females fluctuated on a yearly cycle. The highest percentage of egg-carrying females, i.e. the peak of the reproductive season, lay in the first half of the year. The maximal life span of a generation, from the hatching of the first larvae to the disappearance of the last adults, lasted 1.5 to 2 years. Males moulted into puberty ca one month later than females. The moulting distribution of adult males had roughly the same course as in females. An adult male generation diet out about one to two months before the female generation. Life expectancy was therefore 14 to 17 months for females and 12 to 15 months for males. The sex ratio of juveniles shortly before the pubertal moult was balanced. The sex ratio of adults shifted from 1:1 at the beginning of the reproductive period to ca 1:9 in favour of females at its end.     

Growth of the scyllarid lobsters Ibacus peronii and I. chacei

The growth rates of the morphologically similar scyllarid lobsters Ibacus peronii (Leach, 1815) and I. chacei (Brown and Holthuis, 1998) are described using data from a tag/recapture study and from tagged lobsters kept in captivity. Within particular size classes, we found no differences in moult increments between male and female I. peronii nor between male and female I. chacei. Small individuals of both species always had larger moult increments than larger individuals. For I. peronii, females moulted more frequently than males, and smaller size classes moulted more frequently than larger size classes. Female I. peronii therefore grew more quickly than males and reached their estimated size at sexual maturity (51 mm carapace length) after ∼2 yr. Moulting of I. peronii was seasonal, with most lobsters (96.3%) moulting between October and January. We found no differences in growth rates of I. peronii at two locations along the east coast of Australia: Coffs Harbour in New South Wales (30°18′S; 153°08′E), and Lakes Entrance in Victoria (37°53′S; 148°00′E). For I. chacei, we found no differences in the frequency of moulting between males and females and, because we also found no differences in the moult increments between males and females, the growth rates of both sexes were the same. Received: 14 August 1999 / Accepted: 20 January 2000     

On the timing of moulting processes in reproductively active Northern krill Meganyctiphanes norvegica

The interactions between moult phasing, growth and environmental cues in Northern krill (Meganyctiphanes norvegica) were examined through analysing populations at seasonal, weekly, and daily timescales. The analyses were carried out on resident populations of krill found in three different neritic locations that experience similar environmental signals (the Clyde Sea, Scotland; the Kattegat, Denmark; Gullmarsfjord, Sweden). Seasonal analyses were carried out on the Clyde Sea population and showed that moulting frequency increased significantly moving from winter to summer. The proportion of moulting females in summer samples was often more than double the proportion of moulting males, suggesting that females had a comparatively shorter intermoult period (IMP). Weekly samples taken from the Kattegat showed a similar pattern. However, although the difference between the proportion of female and male moulters was significant in one week, it was not another, mainly because of the variability in the proportion of female moulters. Such variability in females was equally evident in the daily samples taken at Gullmarsfjord. It suggests that females have a shorter IMP (12.5 days) than males (18.4 days) and are more likely to moult in synchrony. Nevertheless, the daily samples revealed that males are also capable of moult synchronisation, although less frequently than females. Shortened IMPs in females were not a result of the abbreviation of specific moult stages. Accordingly, reproductive activity did not alter the course of the normal moult cycle. There was no significant difference between the total body lengths of males and females indicating that females achieve the same levels of growth despite moulting more frequently and having to provision the energy-rich ovaries. This is in contrast to most other crustaceans where the energy costs of reproduction reduce female growth. The fact that females were less abundant than males, probably by suffering a greater level of mortality, suggests that different behavioural strategies, particularly vertical migration regimes, were adopted by each sex to maximise growth and reproduction.     

Costs of overlapping reproduction and moult in passerine birds: an experiment with the pied flycatcher

The process of moult in birds requires the investment of substantial amounts of energy and nutrients in feather production and attendant processes. Flight performance may be reduced during the moult of the wing feathers, and moulting birds may suffer from an increased predation risk. These factors may explain why the moult is usually separated in time from other energetically demanding processes, such as reproduction. In this study, we investigated the importance of the temporal separation of moult from breeding activities in the pied flycatcher Ficedula hypoleuca. We induced a moult-breeding overlap by removing the two innermost primaries on both wings, thus imitating the natural loss of these feathers during the initial stage of the moult. The experiment probably did not stimulate feather regrowth, but may have reduced flight performance. Just before fledging of the young, manipulated males and females, as well as the fledglings, had a lower body condition than control birds. The return rate of adult males in the year following the manipulation was significantly lower for males in the manipulated group than for control males. The return rate of females was lower than that of males, but there was no difference between the two female groups. The number of nestlings recruited in the year after the manipulation was significantly lower for the group with an induced moult-breeding overlap than for the control group. The reproductive performance in recruited and returning old females was independent of the manipulation, and in returning adults and recruits the feather quality in terms of number of barbs/mm was independent of the manipulation. These results suggest that a moult-breeding overlap induces fitness costs in terms of reduced recruitment rate and adult survival in a long-distance migrant. Received: 17 September 1997 / Accepted after revision: 23 February 1998     

Moult in relation to some aspects of reproduction and growth in swarms of Antarctic krill,Euphausia superba

In a 14-d period 38 swarms of Antarctic krill,Euphausia superba Dana, were sampled in an area 55.5 x 55.5 km to the southwest of Elephant Island, at the north of the South Shetland Islands. Moult stage, maturity stage, and size of ca. 100 krill from each swarm were measured. Each of the characteristics varied greatly between swarms. Moulting krill were found in most swarms, but in one swarm all of the krill were just about to moult. Ways in which moulting may act as a possible sorting mechanism are discussed. Data for all the analysed krill (ca. 3000 specimens) were used to investigate the interdependence of moult rate, sexual maturation and growth. While all immature krill moulted at approximately the saine rate in the study, there were significant differences in the moulting rates of mature male and female krill. Gravid female krill continued to moult, although less frequently than mature males. As a consequence males had to attach spermatophores to females after each moult. It is likely that variation in moult rate of females and males was related to the energy expenditure required for ovary development in females and spermatophore production and searching behaviour in males. Spawning and moulting were only partly coupled in gravid females. Spawning appeared to take place predominantly during Moult Stage D₂. There was no evidence of intermoult growth by intersegmental dilation.     

Moulting and reproduction of the spiny lobster Jasus edwardsii (Decapoda: Palinuridae) in northern New Zealand

Aspects of moulting and reproduction of a primarily adult population of the spiny lobster Jasus edwardsii (Hutton) (Palinuridae) were investigated in the field and at an adjacent laboratory in northeast New Zealand from 1982 to 1986. Most males moulted in October/November, ensuring they were in intermoult stage at mating. Females moulted in late April–June, 2 to 3 mo later than females from more southern latitudes. For individual females the specific timing of ecdysis and mating and the length of the interval between them were related to size. Larger females (125 to 135 mm carapace length), tended to moult earlier and wait longer (up to 38 d) before mating, but did so prior to smaller females (95 to 105 mm carapace length, CL) which moulted later, but mated within as few as 9 d. These differences are explained equally well by size-dependent growth processes and by social or chemical inhibition by larger females. Most spiny lobsters (93 to 131 mm CL) moulted at night at depths <10 m. No clear reasons for these behaviours could be found. Eggs were incubated for 101 to 116 d by females in the study population, 70% of the time recorded for more southern areas, these differences probably being related to differences in water temperature. A consequence of earlier female moulting and egg-laying, and longer egg-incubation periods with increasing latitude is that the majority of J. edwardsii larvae hatch in September–November.     

Population structure and growth in captivity of the spiny lobsterPanulirus penicillatus from Dahab,Gulf of Aqaba,Red Sea

Panulirus penicillatus (Olivier, 1791) (Decapoda: Palinuridae) is the most common spiny lobster in the Red Sea and is widely distributed in the Indian and Pacific Oceans. Lobsters (n=377) were collected on four occasions during 1986 on the coral reef at Dahab, Sinai, Egypt. Average size of the collected individuals was 70.5±24.6 mm carapace length (CL) for males and 63.2±15.9 mm CL for females. The sex ratio was 1:1.64 males to females. Length increment per molt was inversely correlated with size and ranged from 2.1 mm per molt in the 40 to 50 mm CL size class to less than 1 mm in the 60 to 70 mm CL size class. Average intermolt period was ca. 136 d for all size classes. The relationship between carapace length and body weight was expressed by the equation:W _b=6.43 × 10^–4 × (CL)^2.89.P. penicillatus from Dahab differ in size, sex ratio and growth rate compared to other palinurid populations throughout their range. This might represent the effect of isolation and location at the edge of the geographical range for this species. It may also indicate an adaptation to their unique habitat in the coral reef in comparison to other palinurid species.     

Life history of the golden ring cowry Cypraea annulus (Mollusca: gastropoda) on Okinawa Island,Japan

Two mark-recapture studies, regular population censuses, field observations, and laboratory culture were used to study the life history of the tropical marine gastropod Cypraea annulus Linnaeus, 1758 from Cape Maeda, Okinawa Island, from April 1984 to March 1986. In the field, the average rate of increase in shell length of marked juvenile snails was 1.0±0.3 mm wk^-1 (N=13), with a maximum of 1.5 mm wk^-1 (initial shell length 10.6 to 17.2 mm). In the laboratory, the maximum growth rate of juveniles was 3.0 mm wk^-1 with food ad libitum. Snails with primordial teeth on the shells grew at a rate of 0.1 to 0.5 mm wk^-1 for about 2 wk. The adults continued to grow at a similar rate for an additional 3 to 6 wk, and ceased detectable growth when some females started spawning egg masses. In the littoral zone on Okinawa Island, snails reproduced throughout the year. Egg masses brooded by three females (shell length 17.4 to 21.3 mm) in the field contained 90 000 to 133 000 ova. In the laboratory, brooding periods of three females lasted 6,8, and 9 d. The estimated average frequency of spawning was 5 egg masses female^-1 yr^-1. The mean shell length of adult females (20.3 mm) was significantly larger than that of adult males (19.6 mm). The life-history strategy of C. annulus is characterized by rapid growth, high fecundity, and repeated spawning throughout the year.     

Reproduction of Halicarcinus planatus (crustacea, decapoda, hymenosomatidae) in the Deseado River estuary, southwestern Atlantic Ocean

The small spider crab Halicarcinus planatus, is a common resident in the rocky coasts of the Magellanic region in South America. The H. planatus population in the Deseado River estuary differs from other populations of the same species in the spatial segregation of sexes. Between March 2002 and October 2003, samples were taken monthly from the lower intertidal and upper subtidal levels. All crabs captured were females. Additional samplings were carried out at the subtidal level to search for the presence of males. A total of 3,346 females were captured and their relative age was determined. Some of these females were maintained in the laboratory to study the terminal moult and the spawning seasons. The analysis of the ovary development of adolescents (ADO) and mature (MAT) females, and the times when females of different ages appear, when spermatophores are found inside spermathecae, and when post-spawning mortality takes place allowed us to establish the whole reproductive process. Adolescents are impregnated only once before the terminal moult. There are seven or eight spawnings after the moult, with a brief resting period between two successive reproductive seasons. The life span of mature females ranges between 15 and 22 months. Results indicate that the H. planatus population of the Deseado River estuary differs greatly from that of the Kerguelen Islands.     

A population study of the polychaete Capitella capitata at plymouth

A population of Capitella capitata (Fabricius) was sampled from January, 1973 to February, 1974. Size of the worms was measured and sexual maturity determined. There is an extensive breeding season, spawning occurring asynchronously throughout the year. Females containing ripe oocytes were almost always present, but most females at any one time contained immature oocytes. A high percentage of males contained active sperm. Oocytes are not released into the coelomic fluid until almost fully developed, and all are released at a single spawning. A large number of eggs are produced at each spawning and there is some evidence to suggest that larval development may be completely benthonic. Gametes first develop in 7 to 8 month old worms and take about 4 months to complete development. Analyses of the female population structure suggests that there are 3 breeding classes. The growth rate is estimated at 30 mm per year, and few worms are believed to survive longer than 2 years.     

Hermaphroditism in brown shrimp: lessons from field data and modelling

This study was undertaken to further clarify whether the brown shrimp, Crangon crangon (Linnaeus 1758), is a gonochorist, a facultative or an obligate hermaphrodite. Juvenile shrimps were sampled from intertidal habitats along the German Wadden Sea coast with a push net and from a power plant water inlet to quantify the share of primary females. Length-based sex ratios were determined for about 27,000 individuals using external characteristics. Observed sex ratios were mainly female-biased, and also large males occurred regularly in the catch. This indicates that sex at hatch is not male as would be characteristic for an obligate protandric hermaphrodite and that not all male shrimps change sex. A cohort-based computer simulation, including sex-specific growth rates, mortality and seasonally varying recruitment, generated sex ratios comparable to the field. The observed decline in the proportion of males with increasing size can be explained solely by faster growth of females without involving hermaphroditism. Based on temperature-dependent growth and moult rates as well as length-specific numbers of eggs per female, the potential egg production of primary and secondary females was modelled, yielding contributions of secondary females of <1%. Sex change in C. crangon has previously been observed and may be interpreted as an evolutionary relict of this species having evolved in a habitat characterized by lower population densities, lower predation levels and increased longevity compared to today’s living conditions in North Sea coastal waters.     

Dynamics of shallow-water populations of Octopus dofleini

Octopus dofleini (Walker) is a common inhabitant of shallow subtidal communities in the Northeast Pacific. The abundances of octopuses at two sites monitored since 1977 have fluctuated greatly during this period. The populations reached high abundances in mid-summer and, during some years, mid-winter. The highest abundances at the two sites did not coincide. There was a constant influx of new octopuses into both study sites, with the greatest immigration occurring in early summer. The octopuses captured spanned a wide range of weights every month, with no clear size classes or seasonal trends in size. The weights of newly-captured octopuses, however, did differ between the sexes and seasons: males weighed more on the average than females, and male weight decreased from winter to fall while female weights did not change O. dofleini appeared to recruit throughout the year; the smallest octopuses occurred between May and November and the greatest number of small octopuses was found in July and August in most years. Females predominated at both study sites throughout the year. However, males predominated among octopuses caught in traps at nearby locations, suggesting that the skewed sex ratios were due to behavioral differences between the sexes.     

Reproductive biology of the spiny lobster, <Emphasis Type="Italic">Panulirus Penicillatus</Emphasis>, in the southeastern coastal waters off Taiwan

The reproductive biology of spiny lobster, Panulirus penicillatus, was studied based on 2,068 lobsters, ranging from 34.28 to 131.60 mm carapace length (CL), sampled in Taitung coastal waters from September 2003 to December 2004. The overall sex ratio approximated 1:1 (χ² = 0.02, P > 0.05), but the monthly sex ratios in 2004 showed significant differences and males were predominant in sizes larger than 80 mm CL. Reproductive activity, assessed using histology, a gonadosomatic index and percentage of ovigerous females, indicated that the mature females could be found in every month and that the major spawning occurred from May to September. The presence of re-developing/re-ripe ovaries by month and size-specific spawning time suggest that larger mature females (>60 mm CL) spawn at least three times a year while smaller new mature females spawn at least once a year. For females, the estimated sizes at 50% physiological and functional maturity were (mean ± SE) 56.46 ± 0.56 mm CL and 66.63 ± 1.07 mm CL. The estimated sizes at functional maturity were between 72 and 74 mm CL for males. The number of eggs per spawning event (brood size, BS) was related to CL by the equation Y _BS = 2.4 × 10^-3CL^4.18 (r ² = 0.902, n = 12). Female lobsters with CL ranging from 60 to 80 mm made the greatest contributions to egg production because of their high brood size and active reproductive activity. A minimum legal size should be established for the fishery to protect egg production potential of lobster population in the southeastern coastal waters off Taiwan.     

Prolonged mating in the milkweed leaf beetle Labidomera clivicollis clivicollis (Coleoptera: Chrysomelidae): a test of the “sperm-loading” hypothesis

Summary Female milkweed leaf beetles (Labidomera clivicollis clivicollis) frequently mate with more than one male, and pairs form mating associations which last for up to 42 h in the field. I tested the hypothesis that males remaining with females for long periods of time benefit by numerically overwhelming the sperm of their competitors. Male L.c.clivicollis copulated intermittently with females throughout an 11 hour period in the laboratory. When virgin females were allowed a single copulation, 94.3% of the sperm they received were located in the spermatheca immediately afterward. Males were not sperm-depleted, for they had large numbers of sperm available after one copulation (mean=230,000±43,200); the maximal number of sperm a male transferred to a female in 24 h was 30,500. There was a positive linear relationship between the number of sperm transferred and time up to 24 h after mounting (r ²=0.178, P<0.003). These data suggest that males transfer increasing numbers of sperm throughout a 24-h-period. Mating duration was the most important determinant of paternity when females were placed with one male for 24 h and another male for 6 hours. Females whose first matings were longer showed first male sperm predominance (as determined by starch-gel electrophoresis), while females whose second matings were longer showed last male sperm predominance. In view of these data, it is puzzling that males do not inseminate with large numbers of sperm immediately after mounting the female. It is possible that female refractory behaviors make insemination difficult and favor prolonged mating by male milkweed leaf beetles.     

Abundance, growth and life cycle of the mesopelagic amphipod Primno abyssalis (Hyperiidea: Phrosinidae) in the Oyashio region, western subarctic Pacific

Abundance and life-cycle features of the mesopelagic hyperiid amphipod Primno abyssalis (formerly P. macropa) in the Oyashio region, western subarctic Pacific, were investigated using samples collected between July 1996 and July 1998. P. abyssalis was collected throughout the entire survey period, with abundance peaks occurring in spring to autumn. While all maturity stages of males and females were observed throughout the study period, the peak reproduction season was in summer. Instar analysis based on the segment number of the pleopod rami indicated that hatched juveniles molted 10 times before becoming adult males and 13 times before becoming adult females. Judging from the dry and ash-free dry weights of each instar, males and females continued to feed throughout the final instar stage. Based on cohort analysis of seasonal samples and laboratory observations on molting frequencies, growth in body length of P. abyssalis was linear with time, and estimated generation lengths were 2.3-3.8 years for females and 1.4-1.9 years for males. Brood size of females ranged from 66 to 337 and increased with increasing female body length. Lifetime fecundity, calculated as the sum of six successive broods, was 1,004. Compared with P. abyssalis in the southern Sea of Japan, those in the Oyashio region have a larger number of adult instars (six versus five for females, three 3 vs one for males), a lower growth rate (0.014 mm day^-1 vs 0.021 mm day^-1), and mature earlier (instar 13 vs instar 15 for females; instar 10 vs instar 11 for males). These characteristics are considered to be advantageous life-history traits to counteract higher niche competition within the mesopelagic community and higher predation pressure by mesopelagic fishes in the Oyashio region than in the Sea of Japan.     

Patterns of sexual cohabitation and female ejaculate storage in the American lobster (<Emphasis Type="Italic">Homarus americanus</Emphasis>)

Little is known of the time and ejaculate allocation strategies during mating of American lobster, Homarus americanus. This study investigated sexual cohabitation and female ejaculate accumulation patterns in a laboratory mating experiment, as well as female seminal receptacle load in exploited populations in the waters of the Magdalen and Anticosti Islands, in eastern Canada. In the laboratory experiment, the length of sexual cohabitation was proportionate to female size for large but not for small males. Also, large males cohabited with pre- and postmolt females longer than small males. These different time investment strategies can be explained by different mutual benefits. In the field and laboratory, larger females accumulated more ejaculate than smaller ones. This suggests that male lobsters tailor ejaculate to female size, a reliable index of her reproductive potential. Moreover, similarly-sized females accumulated more ejaculate when mated with large compared to small males. Comparison of receptacle loads between wild-mated and laboratory-mated females suggests that the former were mating mainly with smaller males, although some evidence of positive size-assortative mating existed, especially at the less exploited Anticosti site. The results are discussed in the context of evolutionary theory and of proposed management measures to increase egg production in exploited populations.Communicated by T. Czeschlik     

Temporal life history plasticity of the Southern Ocean squid <Emphasis Type="Italic">Todarodes filippovae</Emphasis> from waters off Tasmania,Australia

Population dynamics and maturity parameters were analysed for seasonal samples of the oceanic ommastrephid squid Todarodes filippovae from off the coast of Tasmania Australia from 2002 to 2004. Based on assumed daily periodicity in statolith increments, T. filippovae had a life cycle of about a year with the sexually dimorphic females reaching their larger size by predominantly growing faster than males. Due to the small sample size of males, analysis was undertaken on female individuals only. Growth in all samples was best described by a power curve and varied annually, with significantly faster growth in 2001 compared to the subsequent 2 years. Seasonal growth rates also varied with autumn- and winter-hatched squid significantly faster than summer-hatched squid. Spring growth rates were intermediate but not significantly different to the other three seasons. Peak hatching periods occurred in late autumn and early winter. ANOVA revealed a season × year interaction for mantle length and total body weight. Pairwise comparisons showed that the annual differences were likely driven by smaller squid in autumn 2002 compared to autumn samples in the other 2 years. Pairwise comparisons also revealed seasonal differences with winter-caught squid smaller than those from the majority of other samples. There were no seasonal effects on mature female gonad weights but females caught in 2002 had significantly lighter gonad weights than females from the following 2 years. There were no consistent trends among seasons or years in the age structure of mature females. This study revealed the plasticity and flexibility in growth and maturity parameters in this species, with a preference for faster growth during cooler periods.     

Size-assortative pairing across three developmental stages in the Zeus bug, Phoreticovelia disparata

The mechanisms underlying size-assortative pairing have received considerable attention. Typically, pairing is assumed to occur at, or just prior to, the adult phase of the life cycle. However, in many invertebrates, males commence associations with juvenile females who are more than a single moult away from sexual maturity. These species are ideal to explore the importance of reproductive and survival benefits as mechanisms driving size-assortative pairing. In the Zeus bug, Phoreticovelia disparata, adult males are found riding on juvenile (fourth and fifth instar) and adult females—a behaviour that is costly for females but has survival benefits for males. Using a combination of field collections and laboratory manipulations, we show that pairing is size-assortative both within and between female age classes and that riding males are smaller than non-riding males. In a series of mating trials, we revealed that males attempt to ride any female but that their riding success is dependent on female age. We also provide the first direct evidence of female resistance to male riding attempts in P. disparata. We propose that size-assortative pairing arises through adaptations that have evolved to minimise the potential costs of sexual conflict. We suggest that the selective pressure on males to maximise survival benefits is sufficiently high that it outweighs the reproductive benefits of discriminating against fourth instar females. Finally, given that female resistance is under direct selection in juvenile females, it is likely to be the main form of selective pressure for adult females.     

Relationship between reproductive biology and age composition and growth in Urolophus lobatus (Batoidea: Urolophidae)

The aim of the present study was to use samples, collected by trawling throughout the year along the lower west coast of Australia, to determine the reproductive biology, size and age compositions, and growth rate of Urolophus lobatus, a member of a speciose elasmobranch family (Urolophidae) for which there was previously only a small amount of such data. U. lobatus copulates about 3 months prior to the time when females ovulate, which implies that sperm is stored in the oviducal gland of the female in the intervening period. Gestation lasts for 10 months, which is relatively long for a species that does not grow to a large size. This feature is reflected in the fact that the mean disc width at parturition (105 mm) is equivalent to as much as 44% and 52% of the asymptotic disc widths of the females and males of this species, respectively. The trends exhibited by the marginal increments on vertebral centra demonstrated that each of the translucent zones in these centra is laid down annually and that their numbers can thus be used to age this species. Maturity was attained by about 70% of females and 50% of males at the end of the 4th and 3rd years after conception, respectively, or at just over 3 and 2 years after birth, respectively. Females and males typically first reach maturity at ˜200 and ∼160 mm, respectively, and attained maximal disc widths of 277 and 237 mm, respectively. Females reached maximum ages of 15 years from conception and 14 years from birth, with the corresponding values for males being 13 and 12 years, respectively. Since such a large amount of growth occurs during gestation, von Bertalanffy growth curves were constructed using the date of conception, that is, the fertilisation date, as age 0, as is usually the case with teleosts, as well as employing the birth date as age 0, as is typically the case with elasmobranchs. The growth curves drawn through the points for the disc width-at-age of the older female and male fish gave a slightly better fit when using, as age 0, the parturition date rather than the conception date. However, the asymptotic disc widths derived for females and males using the conception date, that is, 241.3 and 202.9 mm, were each still only 7.6 mm less than those derived using the birth date. Moreover, the fit of the line drawn through the points for the disc width-at-age of fish during gestation was better using the conception date as age 0 than would be achieved by a backwards extrapolation of the growth curve using the birth date as age 0. Received: 6 January 2000 / Accepted: 17 June 2000     

Secondary polygyny by inbred wingless sexuals in the dolichoderine ant Technomyrmex albipes

Summary Technomyrmex albipes makes huge polydomous colonies which consist of up to several millions of adults. In field colonies, dealate queens are rare or absent, though winged males and winged females emerge annually (synchronously) in large numbers from late may to mid June. Field and laboratory observations showed that the reproduction of established colonies was performed by wingless females inseminated by wingless males from the same colony. Dissections and morphological examinations revealed that wingless females are workers with no spermatheca and intercastes with a spermatheca. Most intercastes were inseminated, had developed ovaries, and seemed to reproduce, while workers did not seem to reproduce. Extranidal tasks were performed only by workers. Approximately half of the adult population were intercastes, and wingless males represented only a small portion of all adults, the rest being nonreproductive workers. Intercastes and wingless males were produced throughout the year except in winter. The winged females and males copulate outside the nest only after the nuptial flight and the dealate females are able to perform independent founding, but they are also eventually supplanted by intercastes. The adoption of dealate queens by an established natal colony did not seem to occur. Thus we infer that in this species the winged reproductives disperse and found new colonies, while inbred wingless reproductives allow the enlargement and budding of colonies. This species has a special trophic-flow system. There is no trophallaxis among adults, and nutrient transfer from adults to other colony members is achieved exclusively by specialized trophic eggs. All females (dealate queens, intercastes, and workers) seem to produce trophic eggs. This aphid-like life cycle, i.e., the occurrence of both winged and wingless reproductive forms, may have evolved as an adaptation supporting the development of secondary polygyny and polydomy.Offprint requests to: K. Yamauchi