Metabolic rate throughout the annual cycle reveals the demands of an Arctic existence in great cormorants

Aquatic endotherms living in polar regions are faced with a multitude of challenges, including low air and water temperatures and low illumination, especially in winter. Like other endotherms from cold environments, Great Cormorants (Phalacrocorax carbo) living in Arctic waters were hypothesized to respond to these challenges through a combination of high daily rate of energy expenditure (DEE) and high food requirements, which are met by a high rate of catch per unit effort (CPUE). CPUE has previously been shown in Great Cormorants to be the highest of any diving bird. In the present study, we tested this hypothesis by making the first measurements of DEE and foraging activity of Arctic-dwelling Great Cormorants throughout the annual cycle. We demonstrate that, in fact, Great Cormorants have surprisingly low rates of DEE. This low DEE is attributed primarily to very low levels of foraging activity, particularly during winter, when the cormorants spent only 2% of their day submerged. Such a low level of foraging activity can only be sustained through consistently high foraging performance. We demonstrate that Great Cormorants have one of the highest recorded CPUEs for a diving predator; 18.6 g per minute submerged (95% prediction interval 13.0-24.2 g/min) during winter. Temporal variation in CPUE was investigated, and highest CPUE was associated with long days and shallow diving depths. The effect of day length is attributed to seasonal variation in prey abundance. Shallow diving leads to high CPUE because less time is spent swimming between the surface and the benthic zone where foraging occurs. Our study demonstrates the importance of obtaining accurate measurements of physiology and behavior from free-living animals when attempting to understand their ecology.     

Effect of temperature and food availability on reproductive responses of Streblospio benedicti (Polychaeta: Spionidae) with planktotrophic or lecithotrophic development

Streblospio benedicti (Webster) from Tar Landing North Carolina (NC), USA with either planktotrophic or lecithotrophic development were reared under two food levels and three temperature regimes (two mimicking seasonal cycles in NC and one at constant 20°C). During the eight-month experiment no females switched reproductive mode and no significant differences in survivorship or reproductive activity were observed between reproductive types. However, reproductive activity and fecundity-related parameters were subject to influence by food and temperature. Survivorship, body size, and larval production was greater in winter-spring than summer-fall regimes. Higher food levels produced increased survivorship, reproductive activity and egg production in adults with lecithotrophic development but no change in those with planktotrophic development. Body size, egg size, egg number, numbers of larvae per brood pouch, and brood size were strongly correlated in female S. benedicti and most correlation coefficients were similar (or identical) in individuals having planktotrophic and lecithotrophic development. A comparison of egg size and brood size in females from Tar Landing suggests that individuals with the two forms of development package offspring differently but expend approximately equivalent reproductive effort. Larval trophic mode is best viewed as a genetic polymorphism in S. benedicti. Individuals with planktotrophic and lecithotrophic development exhibit similar reproductive responses to environmental variation and there is no evidence for speciation.     

The cost of success: reproductive effort in male southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

Reproductive effort is a key parameter of life history because it measures the resources allocated to reproduction at the expense of growth and maintenance. Male reproductive effort always had a minor role with respect to female effort both in the development of theories and in field research. Elephant seals are an ideal subject for reproductive effort studies because they fast during the breeding season, splitting the phase of energy acquisition from the phase of energy use for breeding. In this paper, we present results on male reproductive effort (weight loss estimated by photogrammetry) in southern elephant seals (Mirounga leonina), the most dimorphic and polygynous of all mammal species. We show that total reproductive effort increases with age, with no sign of late decrease or senescence. Male reproductive effort in this species depends mostly on behavioral factors, i.e., the success in competition with other males, and the intensity of interaction with females. A large effort results in large gains in both mating success and fertilizations. The large reproductive success that a few males are able to achieve come at a big cost in terms of energy expenditure, but this cost does not seem to affect the likelihood of survival to the following breeding season.     

Reproductive effort in Adélie penguins

Summary We estimated reproductive effort (energy expenditures for reproduction, as opposed to maintenance) in Adélie penguins breeding at Palmer Station, Antarctica. Data on body composition changes and metabolic rate were obtained using isotopic methods. Adelie breeding behavior consists of an initial courtship stage (during which both sexes fast), incubation, the guard stage (when chicks are 1 to 18–28 days old), and the creche stage (from the end of guarding until chicks are 28–45 days old). Both males and females lost considerable mass during the initial stages of the reproductive season, but males fasted longer and lost more mass. Mass losses of both sexes consisted of 66% depot fat and 34% lean tissue. Mass and body composition remained constant once birds resumed feeding. The metabolic expenditure for the foraging necessary to accumulate the mass lost while fasting — one component of reproductive effort —was about 63 MJ in males and 39 MJ in females. Field metabolic rates (FMR) were low during courtship and while incubating, increasing more than 2-fold when birds resumed foraging. Although mean FMR increased between incubation and the creche stage, differences between stages were small and not significant. We used FMR data and an energy balance model to estimate the cost of feeding chicks. Results suggest a maintenance FMR of about 2.7 × basal metabolism (BMR), increasing to 3.4–3.6 × BMR during the creche stage. The reproductive effort (as metabolic expenditures) associated with feeding chicks is 31 MJ (males) to 36 MJ (females). Cumulative reproductive effort is 94 MJ in males and 75 MJ in females, or 5.3–6.2% of the annual energy budget. The reproductive effort devoted to chick care does not appear to be constrained by physiological or time limitations. Instead, selection to reduce the risk of predation may prevent the evolution of increased parental care. Correspondence to: M.A. Chappell     

Longevity,calling effort,and metabolic rate in two populations of cricket

Intraspecific variation in a resting metabolic rate (RMR) is likely to be an important determinant of energetic-resource use and may influence the resources subsequently available for allocation to traits not directly associated with somatic maintenance. The influence of RMR on resource availability could be especially important for condition-dependent sexual traits, such as cricket calls, that are themselves energetically costly to produce. RMR may also be associated with longevity, either negatively because individuals with a high RMR burn resources faster and die young, or positively as individuals with high RMR are more able to accrue resources to fuel survival. Additionally, the associations between RMR and other characters may vary across populations if differential selection or drift shapes these traits. Here we tested for differences in RMR, body mass, calling effort, and longevity in two populations of cricket Gryllodes sigillatus and then evaluated the potential influence of RMR on calling and longevity. We find that RMR, calling effort, and longevity varied across populations, but mass did not. Controlling for population and mass, RMR was not significantly associated with calling effort, but was negatively associated with longevity. These findings suggest that male crickets that live fast die young.     

Energy expenditure and food consumption of foraging Imperial cormorants in Patagonia, Argentina

Energy management during the breeding season is crucial for central place foragers since parents need to feed themselves and their offspring while being spatially and temporally constrained. In this work, we used overall dynamic body acceleration as a measure of activity and also to allude to the foraging energy expenditure of breeding Imperial cormorants Phalacrocorax atriceps. We also analyzed how changes in the time or energy allocated to different activities affected the foraging trip energy expenditure and estimated the daily food requirements of the species. Birds spent 42 % of the total energy flying to and from the feeding areas and 16 % floating at sea. The level of activity underwater was almost 1.5 times higher for females than for males. The most expensive diving phase in terms of rate of energy expenditure was descending though the water column. The total foraging trip energy expenditure was particularly sensitive to variation in the amount of time spent flying. During the breeding season, adult cormorants breeding along the Patagonian coast would consume approximately 10,000 tons of food.     

Inter- and intraspecific patterns of reproductive effort in four species of cone shells (Conus spp.)

Patterns of reproductive effort were examined both within and between 4 species of the tropical marine gastropod genus Conus from Hawaii in 1979. Caloric content was measured for the somatic tissues of female cone shells and for their egg capsule masses. These data were used to calculate (1) current reproductive effort, which was the ratio of annual energy produced as spawn to the sum of the annual energy produced as spawn plus the annual energy produced as somatic tissue; and (2) weight-specific reproductive effort, which was the ratio of annual energy produced as spawn to the total energy content of the female body just before spawning. For C. pennaceus, during the approximately 10 yr of life of a female, current reproductive effort rose from about 0.35 at 2 yr to about 0.85 at 10 yr. In contrast, weight-specific reproductive effort remained relatively constant throughout life for C. pennaceus (about 0.2 to 0.5), C. abbreviatus (about 0.2 to 0.3), C. flavidus (about 0.2 to 0.3) and C. quercinus (about 0.1 to 0.2). It is suggested that weight-specific reproductive effort may be useful as an index of the selective importance of the survival cost of present reproduction, while current reproductive effort more closely reflects the selective importance of energy allocation tradeoffs. Data on the 4 species of Conus examined support the hypothesis that reproductive effort should be positively correlated with extrinsic adult mortality and negatively correlated with the variability of juvenile survivorship. The data also suggest a possible relationship between reproductive effort and the cost of parental care.     

Sex-biased maternal expenditure in Rocky Mountain bighorn sheep

Summary We examined maternal energy expenditure in Rocky Mountain bighorn sheep to test the hypothesis (Trivers and Willard 1973) that individual sons in polygynous mammals should obtain a larger maternal subsidy. In accord with theory, males weighed more at birth (an index of relative prenatal expenditure), tended to suckle more between 40 and 100 days of age (an index of relative postnatal expenditure) and imposed greater delays on their mothers' return to estrus in the subsequent breeding season (an index of relative total preweaning expenditure). Mothers rather than offspring appeared to have primary control over postnatal expenditure. The effect of maternal reproductive effort on return to estrus was cumulative over a period of 3–5 years so that ewes making relatively large expenditures gave birth progressively later. Ewe and lamb mortality was associated with late conception in the previous rut. Thus, differential reproductive effort by ewes in this population appeared to entail a fitness cost involving reductions in both offspring and maternal survivorship. Correspondence to: J.T. Hogg at the present address     

Differences in reproductive strategies of morphs of the genus Echinaster (Echinodermata: Asteroidea) from the Eastern Gulf of Mexico

Two morphs of the asteroid genus Echinaster, E. Type 1 and E. Type 2, have reciprocal annual cycles of reproduction and nutrient storage. The pyloric caeca (storage organs) reach maximal size over winter then decrease with increasing gametogenic activity in the spring. E. Type 1 broadcasts buoyant eggs in late May to early June. E. Type 2 deposits benthic eggs in late April. Lecithotrophic larval development is similar in both morphs except that larvae of E. Type 1 undergo a brief planktonic phase. E. Type 1 reaches sexual maturity at a larger size, expends a lower reproductive effort but has a greater absolute reproductive output than E. Type 2. E. Type 2 produces fewer, larger eggs and has a greater parental investment per egg. In both morphs, females have much higher gonadal lipid levels than males and expend a higher reproductive effort in terms organic matter and energy. Reproductive effort, reproductive output, nutrient storage in the pyloric caeca and body size varied between populations and between years in each morph. Egg size and parental investment per egg were constant. Within-morph variability is attributed to differences in nutritional state. Differences in reproductive strategy support the hypothesis that the morphs are separate species.     

Sperm allocation in relation to male traits, female size, and copulation behaviour in freshwater crayfish species

Sperm competition is a well-recognised agent in the evolution of sperm and ejaculate structure, as well as variation in female quality. Models of the evolution of ejaculate expenditure predict that male body condition, female fecundity and the risk and intensity of sperm competition may be the ultimate factors shaping optimal ejaculate size. We investigated sperm allocation in Austropotamobius italicus, a freshwater crayfish exhibiting a coercive mating system and external fertilisation, in relation to male and female traits and copulation behaviour under laboratory conditions. We found that mating males were sensitive to female size and produced larger ejaculates when mating with larger females, which were more fecund in terms of number of eggs produced. We found no evidence for female egg production being sperm-limited, as the number of eggs was not dependent on male sperm expenditure. Copulation duration and number of ejaculations reliably predicted the amount of sperm transferred, and both these behavioural measures positively covaried with female body size. These results indicate that male freshwater crayfish can modulate their sperm expenditure in accordance with cues that indicate female fecundity. In addition, a novel finding that emerged from this study is the decrease in sperm expenditure with male body size, which may either suggest that large, old male crayfish are better able than small males to economise sperm at a given mating to perform multiple matings during a reproductive season, or that they experience senescence of their reproductive performance.     

Contest behavior and other reproductive efforts in aging breeders: a test of residual reproductive value and state-dependent models

Two models have been proposed to explain age-related changes in reproductive performance. State-dependent models predict that reproductive effort depends on the magnitude of surplus energy reserves, which often varies with age. Contrary to this prediction, there was no significant effect of starvation on the outcome of contests for carcasses by female Nicrophorus orbicollis despite weight loss by starved females relative to controls. The residual reproductive value (RRV) model predicts that individuals adjust their current reproductive effort based on potential for future reproduction. Younger adults are predicted to restrain reproductive effort because they are less willing to risk their potentially longer reproductive careers. This model was tested empirically for several components of reproduction. Age was found to be strongly correlated with dominance when two similarly sized females discovered a carcass on the same day. Age also had a small positive effect on egg mass and was positively correlated with ovipositing at least one egg. Age did not affect nesting performance (the degree of carcass burial or the value of a prepared carcass for another female). Age also had no effect on fecundity when a female bred alone but was an important factor when two females were in direct competition. Changing RRV related to aging appears to be a robust determinant of contest outcomes in burying beetles, but effects on other components of reproductive effort in the present and previous studies are more variable, perhaps because of effects of changing state.     

Growth rate of the bathypelagic crustacean Gnathophausia ingens (Mysidacea: Lophogastridae)

The content of water, ash, carbohydrate, lipid, protein, chitin, carbon, hydrogen, nitrogen and energy was measured for all life stages of the bathypelagic mysid Gnathophausia ingens collected in San Clemente Basin off Southern California, USA between January 1969 and February 1982. These data are used to examine the life history, growth rates, growth pattern and partitioning of material and energy over the life of this species. Females reproduce only once and brood their young for about 1.5 yr, during which time the females do not feed. This species has a very high reproductive effort: of the energy accumulated over its life, 61.3% is used in egg production, 13.4% in parental care of young, 5.6% in cast exoskeletons and only 19.6% remains in the females after brooding. The relative rate of growth (percentage of energy d^?1) is about 1% immediately after leaving the mother and declines to about 0.2% as maturity is approached. Such growth rates appear to be typical for an animal of this size living at low temperatures. The evolutionary context of this mysid's life history is discussed.     

Comparative costs of promiscuity and monogramy: A test of reproductive effort theory

Summary Sharp-tailed sparrows Ammospiza caudacuta and seaside sparrows A. maritima co-occupy salt marshes in northeastern North America and are ecologically similar. Female caudacuta care for their young unaided, in comparison to martima's dual parental behavior. The annual reproductive effort of female caudacuta is higher than that of female maritima. Under reproductive effort theory, as both species' reproductive output is the same, female caudacuta should have a lower probability of survival than female maritima. However, the annual survival of both groups was the same. We believe that these results are possible only in highly productive habitats, where the relative cost differences of derived and ancestral mating systems may not be as great as in less productive environments. Lack of penalty in terms of lifetime fitness may have set the stage for the evolution of promiscuity in the sharp-tailed sparrow.In spite of their emancipation from parental duties, male caudacuta survive no better than females. We conclude that activities not directly related to care of young, such as patrolling large home ranges, may entail as much risk or energy expenditure as parental duties do.     

Ovarian development and spawning in relation to the moult cycle in Northern krill, Meganyctiphanes norvegica (Crustacea: Euphausiacea), along a climatic gradient

Adaptive processes linked to reproduction were studied comparatively for three populations of Northern krill, Meganyctiphanes norvegica (M. Sars, 1857), sampled during winter and summer cruises in the Clyde Sea (W Scotland), the Kattegat (E Denmark), and the Ligurian Sea (NW Mediterranean). The aim was to investigate the functional relationship between egg production and moulting under contrasted climatic and environmental conditions. A staging system for female sexual development established for live krill was complemented by a histological study of the ovary at various developmental steps. During the reproductive season, all adult female krill were engaged in cyclical egg production. During experiments, female krill released one batch of mature oocytes in one or two spawning events. The ovary of postspawn female krill still contained developing oocytes for another egg batch. In the non-reproductive period, all female krill had a resting ovary. Ovarian structure and pattern of egg production were identical in the three populations, but seasonal timing of egg production was different. The model proposed for the Ligurian population of the annual cycle of ovarian development can be extended to the other two populations, taking into account the seasonal characteristics of each site. Random field samples were staged simultaneously for moult cycle and for sexual development. Moult stages and the seasonal variation of the intermoult period were studied for the Kattegat population using multi-year data and compared to data obtained during summer/winter cruises in the Clyde and the Ligurian Sea. At the three sites, intermoult period was shorter and temperature-dependent during the reproductive period, concurrent with the season of greatest food availability. During most of the year and the period of sexual rest, moulting activity was reduced. The relationship between spawning and the moult cycle was studied comparatively for the three populations. Eggs were released during the premoult phase of a “spawning moult cycle”, in one or two spawnings associated with apolysis and Moult Stage D1, respectively. Yolk accumulation for the next egg batch was completed during an alternating “vitellogenic moult cycle”. A model for the timing of cyclical egg production in relation to moulting, as proposed for the Kattegat, can be extended to the other populations, taking into account intermoult period variation with temperature. Temperature appeared to be the principal environmental factor controlling growth (through moulting) and egg production during the reproductive season, in connection with favourable trophic conditions. Received: 22 December 1997 / Accepted: 29 August 1998     

Free amino acids as energy substrate in developing eggs and larvae of the cod Gadus morhua

The content of free amino acids (FAA) in the cod (Gadus morhua L.) egg is about 200 nmol at spawning, decreasing by about 100 nmol/egg during the egg stage and about 75 nmol/larva during the yolksac larval stage. Together, alanine, leucine, serine, isoleucine, lysine, and valine account for about 75% of the decrease. Ammonium accumulates gradually during the egg stage and is quickly excreted after hatching. The body protein content is maintained during the egg and yolksac larval stages. The measured oxygen uptake of the cod embryo during the egg and yolksac larval stages accounts for about 85% of the oxygen necessary to catabolize the FAA disappearing during this period. Ammonia excretion of the cod embryo, as taken from literature data, is similar to the expected ammonia production from catabolism of the FAA. Our data suggest that FAA are a major substrate for aerobic energy production in cod eggs and yolksac larvae. The implication of this finding for the production of a favourable first-feed for cod and other cultivated marine fish larvae, and for the selection of high quality eggs of marine fishes, is stressed.     

Pigment allocation to eggs decreases plasma antioxidants in a songbird

Although evidence is accumulating on the adaptive function of female ornamentation, very little is known about maternal allocation decisions involving sexual signaling and other reproductive functions. Blue egg coloration has been suggested as a sexually selected signal of female quality to males, and some recent studies are in accordance with this hypothesis. Blue eggshell coloration results from the deposition of biliverdin pigment by laying females, which is a potent antioxidant. Thus, egg pigmentation should be costly in terms of antioxidants, an assumption of the signaling hypothesis that has not been tested yet. We induced increased reproductive effort in a set of female pied flycatchers Ficedula hypoleuca through nest removal and measured egg pigmentation and plasma antioxidant levels in relation with a control group. Experimental females showed a negative association between egg color and plasma antioxidant levels, while there was no relationship for control birds. This supports that egg pigmentation is costly in terms of general antioxidant defenses and suggests a tradeoff between the allocations to both traits. Furthermore, experimental females with more colorful eggs raised more fledglings, especially when breeding early. Controls did not show a relationship between egg color and reproductive success. Females laying more colorful eggs could have shifted their allocation decisions towards current reproduction, at the expense of their own antioxidant defenses. Our results highlight that blue egg coloration is a life-history trait, subject to tradeoffs with other attributes, and seems to be especially informative under harsh breeding conditions.     

Nest-maintenance effort and health status in chinstrap penguins, Pygoscelis antarctica: the functional significance of stone-provisioning behaviour

Stone provisioning is a nest-maintenance activity accomplished by pygoscelid penguins after reliefs during the incubation/brooding period. The functional significance of this behaviour has been mainly explained as a parental strategy preventing nest flooding under detrimental weather conditions. In addition, and in the light of recent studies, this behaviour could also fit into the sexual selection process. In this study, we tested the first idea, that is, whether stone provisioning is a nest-maintenance behaviour to increase egg/nestling survival by lowering the risk of nest flooding, and can thus be considered a form of parental care. Additionally, we investigated if the effort invested by parents in nest maintenance is constrained by physiologically limiting resources. The effort of stone collection and the perceived risk of nest flooding were experimentally manipulated during the incubation and early brooding phases in a chinstrap penguin, Pygoscelis antarctica, colony. Three groups of nests were established. After weighing, control nests were left unmanipulated. In a second group of nests (reduced group), only one-half of the initial weight of stones was returned to the nests. In a third group of nests (snow-added group), we both reduced nest weight by a half and added snow outside the nest bowl over 6 consecutive days. Ten days after manipulation, the difference in nest weight between initial and final conditions was significantly related to treatment: penguins increased stone provisioning in the reduced group (44% of half-reduced nests), but drastically more in the reduced and snow-added group (123% of half-reduced nests), while the weight of control nests was unchanged compared to premanipulation conditions. The intensity of stone provisioning was affected by nest date, peaking about hatching time and shortly after, and declining with advancing chick age. These results suggest that stone provisioning is a mechanism that has evolved to prevent egg or chick mortality by nest flooding. The haematocrit, but not leukocyte numbers as expressed by the buffy coat layer, varied with the experimental conditions. Penguins investing more time in nest maintenance had a lower haematocrit, suggesting a physiological trade-off probably mediated by competition between the time devoted to nest maintenance versus foraging activities. The amount of stones collected and the haematocrit were positively related to the number of neighbour nests, so those individuals surrounded by more nests seemed to obtain benefits in the availability of nest material and energy savings. This study indicates that stone-provisioning behaviour is a nest-maintenance activity evolved to improve thermal nest characteristics potentially increasing offspring survival, and competing in time and energy with other reproductive activities. Stone provisioning in penguins should therefore be regarded as a form of parental care and an important part of individual reproductive effort in species breeding in harsh environments. Furthermore, nest size and nest-maintenance effort should be considered reproductive traits indicative of parental quality and thus could also be involved in the post-mating sexual selection process.     

Factors affecting the spawning period of sardine in two highly oligotrophic Seas

Ovarian histology and weight dynamics of reproductive and somatic tissues were used to investigate seasonality of spawning and factors affecting reproductive period in two E Mediterranean sardine, Sardina pilchardus, populations (central Aegean and Ionian Seas), between September 1999 and May 2001. Despite similarities in the general spawning pattern of the two populations, i.e. protracted spawning season during the colder months of the year, reproductive period in the Ionian was shorter and lagged behind the Aegean by 2–3 months. In both Seas, reproductive period was influenced by body size as smaller females tended to have both shorter and delayed spawning season. Whereas hepatosomatic index (HSI) was consistently higher in reproductively active females during the whole spawning period, seasonality of spawning did not match variations of HSI. In both Seas fluctuations of HSI matched the spring burst of primary productivity and were followed by fluctuations in somatic condition and mesenteric fat by a 2 months’ lag. Somatic condition and reproductive activity displayed inverse seasonal patterns, implying that egg production in sardine basically relies on visceral and muscular fat deposits, accumulated during the summer growing season. Reliance of egg production on past energetic reserves suggests that sardine is a capital breeder, which is a rather efficient strategy in such oligotrophic habitats as the E Mediterranean, where winter spawning is accompanied by a particular decline in adult preying fields.     

Reproductive biology of the common shrimp Crangon crangon (Decapoda: Crangonidae) in the central Irish Sea

Aspects of the reproductive biology of the common shrimp Crangon crangon (L.) were studied in Port Erin Bay, Isle of Man, Irish Sea. Size at sexual maturity was determined from the proportions of ovigerous females and of females with maturing ovaries. The size at which 50% of females are mature is estimated (±95% confidence intervals) as 12.5±0.48 mm carapace length. Based on the proportions of ovigerous females and of mature females, the main breeding season was from January to June. Mean ovarian dry weights indicated two broods (winter and summer), with females bearing winter broods (WB) having higher gonad indices than those with summer broods (SB). WB females with non-eyed eggs and with eyed eggs differed in the regression of ovarian dry weight on carapace length, indicating preparation for laying a second brood. In both broods the moult stages of berried females were related to egg stage. Moulting will occur following the release of the brood. During embryonic development, mean egg length and egg volume were larger in all stages, and the mean dry weight of individual eggs of all stages heavier, in WB than in SB; there was no difference in egg number, however. Consequently, reproductive investment, the proportion of female weight devoted to egg production, was 67% higher in WB (0.20±0.04) than in SB (0.12±0.03). There is a significant effect of egg volume on brood weight, but not on egg number. In both broods, egg number was a negatively allometric function of female body size in non-eyed eggs and an isometric function of female body size in eyed eggs. Brood mortality during incubation was higher in SB (17%) than WB (10%). Differences in the reproductive variables and investment between the two broods of C. crangon are discussed in the light of reproductive strategies and life history.Communicated by J.P. Thorpe, Port Erin