Comparative life histories of three species of planktonic calanoid copepods in Dabob Bay,Washington

The life history patterns of Calanus marshallae Frost, C. pacificus Brodsky, and Metridia lucens Boeck were determined in Dabob Bay, Washington, USA, during 1973 and October 1981–September 1982. C. marshallae emerged from diapause and moulted to adults in January–February. One major generation was observed, having been produced mainly during early to mid-March. Most surviving individuals spawned in March, arrested development at the C5 stage, and were in diapause by late May. However, some individuals developed to adults by late April and produced a very minor second generation. C. pacificus emerged from diapause and moulted to adults in February–March. The first generation produced during the spring lagged slightly behind the C. marshallae generation. C. pacificus produced two additional generations, one in late spring and one in the fall. The production of the three generations coincided with or closely followed phytoplankton blooms. Some C. pacificus arrested development at the C5 stage and entered diapause during the summer, but the majority of the population did not do so until fall. M. lucens did not appear to enter a diapause state. The fall and winter population was chiefly adult females which mostly remained at depth and were reproductively immature. Nevertheless, naupliar M. lucens were found on all sampling dates, indicating that reproduction never ceased as it did for the two Calanus species. M. lucens produced generations in late winter/early spring, late spring, and late summer/early fall. Interspecific comparisons between the life histories observed in Dabob Bay and intraspecific comparisons between the life histories in Dabob Bay and other locations are made. The advantages or disadvantages conferred to the copepods by their differing life histories are also discussed.     

Annual life cycle of the copepod<Emphasis Type="Italic"> Eucalanus inermis</Emphasis> at a coastal upwelling site off Mejillones (23°S), northern Chile

Eucalanus inermis is an abundant species in the eastern tropical and subtropical South Pacific, including the oceanic and coastal waters off Chile and Peru. Its annual life cycle was studied through a time-series sampling (weekly intervals) during 2002, at a fixed coastal station at an upwelling site (Mejillones Bay, 23°S) off northern Chile. The more-or-less continuous occurrence and abundance of naupliar and copepodid stages indicated that the species reproduces during most of the year, with two peaks: one during the austral spring and the other during the summer. Thereafter, an abrupt decline in the population was observed during winter. The abundances of E. inermis copepodids and nauplii were positively correlated with sea surface temperature, suggesting temperature-dependent development and growth during the spring–summer period. Three cohorts could be distinguished during one annual cycle, with generation times >30 days. The estimate of mean weight-specific daily growth (0.12 day^–1) is lower than that of other species in the area, but it is consistent with a slower development rate. The distribution of this copepod is associated with the equatorial subsurface waters characterized by low-oxygen content (<0.5 ml l^–1). The ascent of this water mass to the near-surface during coastal upwelling in the spring–summer period and the presence of E. inermis adults favor their reproduction and the development of cohorts in the food-rich upwelling environment. The wintertime disappearance of E. inermis from shallow waters is, thus, interpreted as a movement to deep waters (>200 m depth), probably in a lethargic mode, within the oxygen minimum zone in the adjacent oceanic area.Communicated by P.W. Sammarco, Chauvin     

Seasonal changes in abundance and development of<Emphasis Type="Italic"> Calanus pacificus</Emphasis> (Crustacea: Copepoda) in the Oyashio–Kuroshio Mixed Region

Seasonal changes in abundance and development of Calanus pacificus Brodsky were investigated by analyzing samples of different depth strata (0–150 m and 0–1000 m) collected monthly in the Oyashio–Kuroshio Mixed Region. Copepodite stage 5 (C5) emerged from dormancy and matured in early summer. A new generation appeared in July and developed into C5 during summer to autumn. Some of the summer generation arrested development at C5 and persisted below 150 m depth until the following early summer. Although the remainder matured and reproduced in October, a new generation was not observed at the surface during winter. These results suggest that C. pacificus shows two different life-cycle patterns, i.e. one generation annually, with overwintering C5 in deep waters, and two generations annually, with surface development during autumn to winter. The complex life-cycle patterns may be an adaptation to the highly fluctuating surface environment in the Oyashio–Kuroshio Mixed Region.Communicated by T. Ikeda, Hakodate     

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.     

The amphipod Corophium multisetosum (Corophiidae) in Ria de Aveiro (NW Portugal). I. Life history and aspects of reproductive biology

The population of Corophium multisetosum Stock, 1952 in Areão displayed a semiannual, iteroparous life history. Mean longevity was ～6?mo, with the estimated life span longer for overwintering individuals born in autumn than for individuals born in spring. Length-frequency data indicated that the length increment per moult is probably higher in males than females; however females moulted more frequently and achieved a larger body size. Preliminary growth rates were 100?μm?d^?1 for juveniles and 19 to 29?μm?d^?1 for mature females, with the lower values occurring during the winter. It was estimated that under favourable conditions females may attain reproductive size and mature within 1?mo. Although incubating females were present all year round, recruitment occurred in spring, almost ceased during the summer, peaked in autumn, and decreased again during the winter. Extreme temperatures and very low salinities during winter and summer may have deterred breeding, while moderate temperatures (15 to 20?°C) and salinities > 1?psu in spring and autumn were apparently favourable for reproduction. The unfavourable summer conditions constrained breeding and synchronised the timing of reproduction. In late-autumn and during the winter, as temperature decreased and brooding time increased, synchrony was progressively lost. Brood size varied as a function of embryonic developmental stage, size of incubating females, and season. The life-history pattern and reproductive features of C. multisetosum in Areão are closely related to temperature and salinity; other environmental conditions such as oxygen content of the water and food availability may also be relevant.     

Male backspace availability in the giant waterbug (Belostoma flumineum Say)

Summary Following bouts of courtship and copulation, female giant waterbugs (Belostoma flumineum Say) deposit eggs on the backs of their mates. Throughout a 6– to 12-day brooding period, males display several behaviors that are vital to egg-nymph survival. Consequently, females depend on male post-copulatory behaviors for successful reproduction and the possibility exists for male backspace availability to limit female reproduction in this species. I studied seasonal trends and factors that affect male backspace availability in populations of B. flumineum in east-central Illinois (USA). Early in the spring/summer, giant waterbug populations are relatively small and a large majority (188/205=91.7%) of the males are egg-laden; males experimentally added to the population during this period quickly became encumbered. In contrast, later in the summer after young-of-the-year emerge as adults, the waterbug population density increases dramatically and fewer (670/1274\2= 52.6%) of the males are encumbered (egg-laden). Of the males that are egg-laden both early and late in the season, significantly more are completely encumbered (i.e., 100% of the dorsum egg covered) early in the spring. The adult sex ratio is generally not biased and the number of eggs/pad that completely covers a male approximates a full ovarian complement. Therefore, these factors probably do not cause male backspace to become limited. The primary factor that appears to limit male backspace availability is the ability of females to synthesize a second partial clutch in a short time, often within 1 to 4 days. Females are capable of ovipositing partial clutches on 12 males within 30 days, whereas male brooding period is temperature dependent and ranges from 6 to 13 days. Newly emerged males are capable of breeding significantly sooner than can newly emerged females, thereby creating ample oviposition substrate for the females in the population after young-of-the-year adults appear. The evolution of sex-role reversal is not well understood; however it should not evolve in waterbugs unless male backspace limits female reproduction. Such a situation appears to exist in B. flumineum early in the season but not later in the summer.     

Egg production and feeding by copepods prior to the spring bloom of phytoplankton in Fram Strait,Greenland Sea

Reproductively activeCalanus hyperboreus (Krøyer) andC. glacialis Jaschnov were captured in the upper 100 m of Fram Strait (77° to 79°N) in late winter 1987. There was no evidence of a phytoplankton bloom; chlorophylla concentrations were uniformly low (<0.1 mg m^–3), and nitrate concentrations were uniformly high (>11.3 mg-at m^–3). Gut-fullness measurements indicated that females were ingesting very little. The maturation state of gonads of bothC. hyperboreus andC. glacialis indicated that 75% of females were in a ripe condition consistent with observed egg laying. The lipid content of females laying eggs was reduced in both species compared to that of females not laying eggs. InC. hyperboreus the reduction was 39% and inC. glacialis it was 44%. All the evidence suggests that bothC. hyperboreus andC. glacialis were laying eggs in late winter by using lipids stored previously; they were not relying on ambient concentrations of phytoplankton. The daily rate of egg laying byC. glacialis using lipids in late winter exceeded the rate reported for summer when ambient food supplies have been shown to be necessary. It is suggested that individuals, spawned well in advance of the spring bloom of phytoplankton, may comprise a major portion of the annual recruitment to the entire population ofC. glacialis in this area, and that their life cycle can be completed within 1 yr. NeitherMetridia longa (Lubbock) norC. finmarchicus (Gunnerus) laid eggs during this study.     

Biologie d'un copépode des mares temporaires du littoral méditerranéen français: Eurytemora velox

The reproduction of Eurytemora velox, a brackish copepod from temporary lakes of the south of France, was studied in winter and spring 1978, under various temperature and salinity conditions, using Chlorella sp. and Amphidinium sp. as food. Maximum numbers of successive eggs sacs (9), eggs per sac (39.3) and total egg production per female (311) were recorded for the witner generation, only 4.8, 34.8 and 109, respectively, for the spring generation. In contrast, the number of eggs per female per day was highest (11.3) in the spring generation, which displayed a more rapid reproductive cycle. Despite strong individual variations in the rhythm of egg sac production and in abundance of eggs per sac, egg production was generally higher during the first third of adult life, attaining a maximum after production of the second or the third egg sac. The continuous presence of the males was necessary to assure complete fertilization of eggs throughout the whole life of adult females. Hatching rate displayed high individual variability, in particular for the spring generation, which had lower average hatching rates (between 0 and 26%, depending on salinity or temperature) than the winter generation (14 to 64%). These differences may be related to the ability of E. velox to produce resting eggs during spring, allowing the species to maintain itself in a temporary water milieu. Temperature significantly affected longevity and daily egg production of females; presence or absence of males did not affect these parameters. An increase in salinity from 20 to 30%. reduced longevity, number of egg sacs, and daily egg production in the winter generation, but not in the spring generation. The specific daily production of females during their adult life was calculated from the egg production:biomass ratio of females, in carbon units. In the winter generation, this ratio increased between 10° and 15°C compared to ratios between 15° and 20°C; the opposite was observed for the spring generation. The seasonal differences in the effects of temperature and salinity on reproduction could indicate an adaptation mechanism to the strongly thermal and haline seasonal fluctuations which characterize the habitat of Ex. velox (brackish waters, drying-up in summer). Larval mortality was high, except at 20%. S for the spring generation. The sex-ratio of the offspring was unaffected by variations in breeding conditions. Hatching time and development time of larvae could be described by two Blehrádek equations displaying close b and () coefficients. We calculated the energy balance of adult females from data obtained in a previous study on feeding and respiration in E. velox, and this is discussed in context with the egg production results. Net growth efficiency varied with algal concentration according to an asymptotic curve, reaching a maximum of 0.43 with Tetraselmis maculata as food or 0.53 with Amphidinium sp. Actual egg production rate obtained in the present study was in good agreement with that calculated by the difference between assimilated food and respiration expenses.     

Seasonal variations in distribution patterns of sympagic meiofauna in Arctic pack ice

During two expeditions of the R.V. Polarstern to the Arctic Ocean, pack ice and under-ice water samples were collected during two different seasons: late summer (September 2002) and late winter (March/April 2003). Physical and biological properties of the ice were investigated to explain seasonal differences in species composition, abundance and distribution patterns of sympagic meiofauna (in this case: heterotrophs >20 µm). In winter, the ice near the surface was characterized by extreme physical conditions (minimum ice temperature: –22°C, maximum brine salinity: 223, brine volume: 5%) and more moderate conditions in summer (minimum ice temperature: –5.6°C, maximum brine salinity: 94, most brine volumes: 5%). Conditions in the lowermost part of the ice did not differ to a high degree between summer and winter. Chlorophyll a concentrations (chl a) showed significant differences between summer and winter: during winter, concentrations were mostly <1.0 µg chl a l^–1, while chl a concentrations of up to 67.4 µmol l^–1 were measured during summer. The median of depth-integrated chl a concentration in summer was significantly higher than in winter. Integrated abundances of sympagic meiofauna were within the same range for both seasons and varied between 0.6 and 34.1×10³ organisms m^–2 in summer and between 3.7 and 24.8×10³ organisms m^–2 in winter. With regard to species composition, a comparison between the two seasons showed distinct differences: while copepods (42.7%) and rotifers (33.4%) were the most abundant sea-ice meiofaunal taxa during summer, copepod nauplii dominated the community, comprising 92.9% of the fauna, in winter. Low species abundances were found in the under-ice water, indicating that overwintering of the other sympagic organisms did not take place there, either. Therefore, their survival strategy over the polar winter remains unclear.Communicated by O. Kinne, Oldendorf/Luhe     

Studies on the growth,reproduction, and life cycle of the supralittoral isopod Ligia pallasii

A two-year study on a field population of Ligia pallasii Brandt has shown that the isopods live for 1.5 to 2 years. Breeding occurs in the spring and early summer, with some females carrying winter broods of eggs. The mean length of breeding females is 22.5±2.2 mm (standard deviation) and the mean brood size is 48±11 eggs. Mature males are larger than mature females (900 and 300 mg live weight, respectively), and are disproportionately broader (31×17 mm and 22.5×9 mm, respectively). The larger size and breadth of the males is an adaptation for copulation, and may be atributed in some measure to slower growth of the female due to the extra energy demands of reproduction. The overall 1:1 ratio of males to females in the population represents the balance between an equal sex ratio in the immature stages, more females in the 18 to 24 mm length category, and more males in the larger length categories. This condition is attributed mainly to the faster growth of the males.     

Comparative life-history study on sympatric hyperiid amphipods (<Emphasis Type="Italic">Themisto pacifica</Emphasis> and <Emphasis Type="Italic">T</Emphasis>. <Emphasis Type="Italic">japonica</Emphasis>) in the Oyashio region,western North Pacific

Life-history features of the sympatric amphipods Themisto pacifica and T. japonica in the western North Pacific were analyzed based on seasonal field samples collected from July 1996 through July 1998, and data from laboratory rearing experiments. T. pacfica occurred throughout the year, with populations peaking from spring to summer. In contrast, T. japonica were rare from autumn to early winter, but became abundant in late winter to spring. Mature T. pacifica females and juveniles occurred together throughout the year, indicating year-round reproduction. Mature T. japonica females were observed only in spring, and juveniles occurred irregularly in small numbers, suggesting limited, early-spring reproduction in this study area. Size composition analysis of T. pacifica identified a total of eight cohorts over the 2 years of the study. Due to the smaller sample size and rarity of mature females (>9.6 mm) and males (>7.1 mm), cohort analyses of T. japonica were not comparable. Laboratory rearing of specimens at 2°C, 5°C, 8°C and 12°C revealed that a linear equation best expressed body length growth by T. pacifica, while a logistic equation best expressed body length growth by T. japoncia. Combining these laboratory-derived growth patterns with maturity sizes of wild specimens, the minimum and maximum generation times of females at a temperature range of 2–12°C were computed as 32 days (12°C) and 224 days (2°C), respectively, for T. pacifica, and 66 days (12°C) and 358 days (2°C), respectively, for T. japonica. The numbers of eggs or juveniles in females marsupia increased with female body length and ranged from 23 to 64 for T. pacifica and from 152 to 601 for T. japonica. Taking into account the number of mature female instars, lifetime fecundities were estimated as 342 eggs for T. pacifica and 1195 eggs for T. japonica. Possible mechanisms for the coexistence of these two amphipods in the Oyashio region are also discussed.Communicated by O. Kinne, Oldendorf/Luhe     

Reproductive strategies and life histories in the cheilostome marine bryozoans Chartella papyracea and Bugula flabellata

Seasonal life history phenomena were monitored through 1978 for two sublittoral cheilostome bryozoans from souther Britain: Chartella papyracea (Ellis and Solander), a non-placental brooder, and Bugula flabellata (Thompson in Gray), which is placental. The intracolonial relationships between growth by zooid budding, polypide recycling, and sexual reproduction, were analysed for both species — quantitatively in the case of C. papyracea. Although both species grow only in summer, their life histories differ. C. papyracea is more of a K-strategist, producing perennial fronds that release larvae in several successive years, most being liberated in winter months. The reduced need for maximization of the larval productivity rate is reflected in the intracolonial configuration of polypide and sexual recycling. B. flabellata is more of an r-strategist. The larvae are produced by the ephemeral fronds of two successive colony generations each summer. Colonies are dormant in winter. The need to maximize the larval productivity rate in this species is reflected in the polypide/sexual recycling configuration. It is believed that embryonic placentation contributes to the maximization of the larval productivity rate in this and other placental brooders.     

Cycle biologique de l'isopode Jaera hopeana dans les milieux marins et lagunaires du Nord-Est de la Tunisie

A comparative study shows that invasion of some salt-water lakes by Jaera hopeana Costa, 1853, a commensal species of Sphaeroma serratum, has not induced any notable modifications in its life cycle. This life cycle has been defined by following mensual evolution in the composition of field populations. Reproduction occurs throughout the year, but is more intense during some periods; growth is continuous in immature individuals, but ceases in summer in males and females. The life span is about 6 to 10 months; the species has, therefore, a short generation time.     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

The reproductive cycle of the Thecosomatous pteropod Limacina retroversa in the western South Atlantic

The reproductive cycle of Limacina retroversa in the Argentine Sea was studied by gonadal analysis. Samples were collected in 1978–1979. The relation between gonadal maturity and size is dependent on the season and the geographical area. Evidence of in situ reproduction was found at the beginning of spring and at the end of summer, indicating that the life cycle in the area consists of two generations per year. In accordance with the season of birth, both generations develop different strategies. Individuals born in spring mature early and reproduce before the end of summer. The offspring born in summer survive the cold season without reaching sexual maturity and reproduce in the following spring.     

Metabolism and body composition of a copepod (<Emphasis Type="Italic">Neocalanus cristatus</Emphasis>: Crustacea) from the bathypelagic zone of the Oyashio region,western subarctic Pacific

Metabolism [respiratory oxygen consumption, electron-transfer-system (ETS) activity] and body composition [water, ash, carbon (C), nitrogen (N), carbon/nitrogen (C/N) ratio] of stage C5/C6 Neocalanus cristatus from 1000 to 2000 m depth of the Oyashio region, western subarctic Pacific, were determined during the period of July 2000 through June 2003. Compared with the C5 specimens from shallow depths (<250 m), those from 1000 to 2000 m were characterized by quiescent behavior, reduced respiration rates (30% of the rates at active feeding), very low water content (61–70% of wet weight), but high C content (56–64% of dry weight) and C/N ratios (7.2–10.6, by weight). Artifacts due to the recovery of live specimens from the bathypelagic zone appeared to be unlikely in this study, as judged by the consistent results between re-compression (100 atm) and non-compression (1 atm) respiration experiments, and between ETS activities and respiration rates directly measured. In addition, the respiration rates of C6 males and females of N. cristatus from the same 1000–2000 m depth were two to three times higher than the rates of C5 individuals, but were similar to the rates of a bathypelagic copepod, Paraeuchaeta rubra. Combining these results with literature data, C budgets of: (1) diapausing C5 specimens, weighing 6–10 mg dry weight; (2) molt to C6 females; and (3) the complete the life span were established, taking into account assorted losses in respiration during diapause at stages C5 and C6, molt production and egg production. Respiratory C losses by C5 and C6 specimens were estimated on the basis of body N as adjusted metabolic rates [AMR; µl O₂ (mg body N)^–0.843 h^–1], then N budgets were also computed subtracting N lost in the form of cast molts and eggs from the initial stock. Calculations revealed that allocation of the C stock was greatest to egg production (34–57%), followed by respiration (27%) and cast molts (3%), leaving residual C of 13–36% in spent C6 females. The present results for N. cristatus from the North Pacific are compared with those of Calanus spp. in the North Atlantic.Communicated by O. Kinne, Oldendorf/Luhe     

Infant care,infanticide, and female reproductive strategies in polygynous groups of common marmosets (Callithrix jacchus)

Reproduction in marmoset and tamarin groups is typically restricted to a single dominant female, but it is unclear why subordinate females tolerate delayed reproduction. The presence of two breeding females in free-ranging groups of common marmosets (Callithrix jacchus) presented a unique opportunity to examine differences in the reproductive strategies of dominant and subordinate females. Three groups were monitored for 12–18 months at a forest reserve in northeastern Brazil. Data on infant care were collected during two consecutive all-day follows every 10 days until the infants were 2 months old. Carrying patterns for infants born to dominant females were similar to those observed in groups containing a single breeding female. All group members over 5 months of age participated in infant care, and dominant females allowed some group members to carry their infants from the 1 st day of life. In contrast, subordinate females were protective of their offspring and were their sole caretakers for at least a week following birth. One infant born to a subordinate was killed in an attack involving the dominant female. Overall, dominant females gave birth to more infants and had higher infant survival than did subordinate breeding females. subordinate females were successful in rearing young only when the timing of births was such that they did not overlap with the dependency period of infants born to the dominant female. These patterns suggest that subordinate females may face resource competition, especially over access to helpers. In times of severe competition, subordinate females that delay reproduction may be avoiding a wasted reproductive effort.     

Life history of the sandy-beach amphipod Neohaustorius schmitzi (Crustacea: Haustoriidae)

Neohaustorius schmitzi Bousfield, the dominant macroscopic invertebrate on sandy beaches in North Carolina, USA, is strictly intertidal, and distributed from mean low tide to high tide, with an average density of 790/m². Location of greatest densities within the intertidal zone is dependent upon season and sex. An aggregated distribution is shown. N. schmitzi has 2 generations/year: a winter generation which lives about 8 months, and a summer generation which lives about 4 months. The reproductive season extends from February through October, with spring and summer peaks. Each female appears to produce only 1 brood of young. The number of young ranges from 2 to 14, depending upon the length and generation of the female.     

Life-history analysis in “physiological” compared with “sidereal” time: An example with an amphipod (Gammarus lawrencianus) in a varying environment

The multivoltine, estuarine amphipodGammarus lawrencianus has four generations per year in an environment where temperatures range seasonally from –1° to 25°C. Temperature-response curves for rates of brood production and development were determined by laboratory experiments and field observation. The life history and population dynamics were observed over a full annual cycle (1981) for a field population located at Rocky Run, Porter's Lake, Nova Scotia, Canada. On a natural (i.e., sidereal) time scale, the generations appear to have very different life histories: the two summer generations have short lives, rapid development and mature at small size (classicr-selection), whereas the overwintering generations have relatively low rates of mortality, slow development and mature at large size (classicK-selection). This pattern (larger size at maturity at lower temperatures) is widespread in aquatic poikilotherms. Similar life-history differences are evident among cohorts of the summer generations that mature at different temperatures. When time is expressed on a physiological scale that removes the effect of temperature on embryonic development and reproductive rate, the variation within and among generations is greatly reduced. In particular, an apparent alternation betweenr- andK-selection largely disappears. Because the generations are temporally isolated, it might be surmised that natural selection acting on the summer generations might antagonize the effects of natural selection acting on the fall and winter generations. However, the scaling of the rates of development, maturation, growth, reproduction and mortality on the physiological time scale derived from the temperature dependence of development and reproductive rate gives a very different and more homogeneous pattern.     

Spawning pattern and type of fecundity in relation to ovarian allometry in the round herring <Emphasis Type="Italic">Etrumeus teres</Emphasis>

Spawning pattern (assessed by seasonal changes in ovarian developmental stages) and type of fecundity (assessed by analysis of oocyte-size frequency distributions) of the round herring Etrumeus teres were studied in relation to ovarian growth and seasonal changes in the gonadosomatic (GSI), hepatosomatic (HSI) and liposomatic (LSI) index as well as the somatic condition of spawners (CS) in a spawning ground of southern Japan. Except for summer, mature and recently spawned ovaries occurred all year round. Oogonia and primary oocytes were present in all ovaries, and cortical alveoli stage (CA) oocytes occurred in all mature, hydrated and partially spent (PS) females (PS: females containing post-ovulatory follicles). Before hydration, a clutch of larger yolked oocytes, undergoing synchronous growth (range 0.7–1.1 mm), was present in mature ovaries which was completely separated from a more heterogeneous clutch of oogonia, primary and secondary oocytes (<0.150 mm) and oocytes in the CA stage (range 0.15–0.60 mm). As vitellogenesis progressed, the yolked clutch increased in size but the CA oocytes remained arrested. The latter entered into the secondary growth phase when hydration started in the advanced batch. Ovarian growth was isometric in all developmental stages, validating the use of GSI, which showed a consistent monthly evolution among years. Spawning stopped in summer (July and August) and peaked in winter and spring. HSI correlated positively with GSI on both a monthly mean basis (r = 0.76) and individual fish basis (liver weight explained 67–83% of the variability in ovary weight when females were grouped into 1-unit GSI intervals) suggesting a significant role of liver in vitellogenesis. LSI and CS also showed marked seasonal changes peaking from summer to middle autumn. Overall results suggest that E. teres is a multiple spawner with a group-synchronous ovarian development and indeterminate annual fecundity, with the three processes linked to an isometric growth of the ovary. We propose that such a reproductive pattern is an adaptation to produce batches of large pelagic eggs through a protracted spawning season.