Aggregation of whelks, Buccinum undatum, near feeding predators: the role of reproductive requirements

In the Mingan Islands, northern Gulf of St Lawrence (eastern Canada), the whelk Buccinum undatum displays a strong escape response to its predator, the asteroid Leptasterias polaris, nevertheless large sexually mature individuals occasionally approach feeding L. polaris to obtain food. In this study, we investigated the hypothesis that reproductive requirements increase the tendency of sexually mature whelks to approach feeding asteroids. Prior to egg laying, females (which invest more energy than males into the production of reproductive structures) represented 72% of the adult whelks that approached feeding L. polaris, but only 36% of the adults randomly collected from the study area. Furthermore, females that were attracted to feeding asteroids had smaller reproductive organs (after accounting for body size) than females randomly collected from the study area. Similarly, prior to egg laying, females fed longer and ingested more food than males when tested in the presence of L. polaris in the laboratory. After egg laying, however, females and males displayed a similar tendency to feed in the presence of a predator, both in the field and in the laboratory. Predator-impact indices, computed by contrasting the feeding activity of whelks in the absence and presence of a predator, indicated that females (but not males) responded more boldly to predators prior to than after egg laying, despite a general decrease in feeding activity at that period. Taken together, our observations indicate that the tendency of adult whelks to approach feeding predators is influenced by potential reproductive gains. Because such gains are presumably more directly linked to a given feeding opportunity in sexually active individuals, whelks may be selected to display increased levels of boldness towards predators with the onset of sexual maturity. Thus, potential reproductive benefits may partly explain the size-dependent tendency of whelks to approach feeding asteroids. Copyright 2001 The Association for the Study of Animal Behaviour.     

Nature and role of newly described symbiotic associations between a sea anemone and gastropods at bathyal depths in the NW Atlantic

The hormathiid sea anemone Allantactis parasitica was found living as an epibiont on numerous species of gastropods at depths of 725-1100 m along the continental slope of eastern Canada. The proportion of bathyal gastropods hosting 1-6 sea anemones reached 72.5% in a single trawl. Although A. parasitica was occasionally found on other substrata (i.e. empty shells, pebbles), laboratory trials confirmed that they preferably associate with living gastropods. Settlement of planula larvae occurred significantly more often on the shells of live bathyal gastropods than on all other substrata present in the tanks. Juvenile sea anemones (∼ 1 mm diameter) readily moved from the mud or other inert substrata onto shells of burrowed bathyal gastropods. Conversely, larvae, juveniles and adults of A. parasitica never associated with any shallow-water gastropods when given the opportunity. Trials exposing predatory sea stars (Leptasterias polaris) from shallow and bathyal depths to bathyal gastropods (Buccinum undatum) with epibiotic A. parasitica, and to asymbiotic bathyal and shallow-water B. undatum, revealed adaptive behaviours in both prey and predator. Shallow-water gastropods (devoid of epibionts) reacted defensively to L. polaris, whereas bathyal gastropods relied mostly on their epibionts to protect them, thus falling prey to L. polaris when the epibionts were removed. L. polaris from bathyal depths typically ignored symbiotic gastropods, but they consistently preyed on asymbiotic ones, while L. polaris from shallow areas initially attempted to prey on all gastropods, but learned to avoid those harbouring sea anemones. Furthermore, living as epibionts afforded sea anemones a means to escape one of their own predators, the sea star Crossaster papposus. The mutualistic relationship between hormathiid sea anemones and bathyal gastropods from the NW Atlantic may have evolved in response to predation pressure.     

Context-Dependent Plastic Response during Egg-Laying in a Widespread Newt Species

Previous research on predator-induced phenotypic plasticity mostly focused on responses in morphology, developmental time and/or behaviour during early life stages, but the potential significance of anticipatory parental responses has been investigated less often. In this study I examined behavioural and maternal responses of gravid female smooth newts, Lissotriton vulgaris, in the presence of chemical cues originating from invertebrate predators, Acilius sulcatus water beetles and Aeshna cyanea dragonfly larvae. More specifically, I tested the extent of oviposition preference, plasticity in egg-wrapping behaviour and plasticity in egg size when females had the possibility to lay eggs at oviposition sites with and without predator cues during overnight trials. I found that individuals did not avoid laying eggs in the environment with predator cues; however, individuals that deposited eggs into both environments adjusted the size of the laid eggs to the perceived environment. Females deposited larger eggs earlier in the season but egg size decreased with time in the absence of predator cues, whereas individuals laid eggs of average size throughout the investigated reproductive period when such cues were present. Also, egg size was found to be positively related to hatching success. Individuals did not adjust their wrapping behaviour to the presence of predator cues, but females differed in the extent of egg-wrapping between ponds. Females’ body mass and tail depth were also different between ponds, whereas their body size was positively associated with egg size. According to these results, female smooth newts have the potential to exhibit activational plasticity and invest differently into eggs depending on temporal and environmental factors. Such an anticipatory response may contribute to the success of this caudate species under a wide range of predator regimes at its natural breeding habitats.     

Dietary carotenoid supplementation facilitates egg laying in a wild passerine

During egg laying, females face a trade‐off between self‐maintenance and investment into current reproduction, since providing eggs with resources is energetically demanding, in particular if females lay one egg per day. However, the costs of egg laying not only relate to energetic requirements, but also depend on the availability of specific resources that are vital for egg production and embryonic development. One of these compounds are carotenoids, pigments with immuno‐stimulatory properties, which are crucial during embryonic development. In this study, we explore how carotenoid availability alleviates this trade‐off and facilitates egg laying in a small bird species, the blue tit (Cyanistes caeruleus). Blue tits have among the largest clutch size of all European passerines and they usually lay one egg per day, although laying interruptions are frequent. We performed a lutein supplementation experiment and measured potential consequences for egg laying capacity and egg quality. We found that lutein‐supplemented females had less laying interruptions and thus completed their clutch faster than control females. No effects of treatment were found on the onset of egg laying or clutch size. Experimentally enhanced carotenoid availability did not elevate yolk carotenoid levels or egg mass, but negatively affected eggshell thickness. Our results provide hence evidence on the limiting role of carotenoids during egg laying. However, the benefits of laying faster following lutein supplementation were counterbalanced by a lower accumulation of calcium in the eggshell. Thus, even though single components may constrain egg laying, it is the combined availability of a range of different resources which ultimately determines egg quality and thus embryonic development.     

Alterations in serum steroid concentrations in the clearnose skate, Raja eglanteria: correlations with season and reproductive status.

Serum steroid hormones in the peripheral circulation of the clearnose skate, Raja eglanteria, were measured at the time of capture and at various times throughout the year while the animals were maintained as a captive breeding population. These analyses demonstrate interesting correlations between changes in hormone concentrations and annual reproductive events. Animals were sampled once (78 females, 20 males) or multiple times (15 females). For both groups of females, 17beta-estradiol was detected throughout the year with significant elevations occurring during October and November when ovarian follicles begin to mature (as determined through necropsy examinations), and January and February when maximum mating activity is observed and egg laying begins. Testosterone and dihydrotestosterone concentrations were significantly elevated in females only during January and February. Testosterone elevations were synchronous with longer-term elevations in 17beta-estradiol in females sampled either once or repetitively. Testosterone concentrations in males were significantly elevated during times of maximum breeding activity compared to periods of sexual inactivity. Data from females sampled during five stages of the egg laying process, as defined by the position of palpable egg capsules within the reproductive tract, revealed that 17beta-estradiol was highest when egg capsules were forming in the nidamental gland (stage 2) or uterus (stage 3); testosterone and dihydrotestosterone were maximal when eggs were in the uterus (stage 3) or cloaca (stage 4); and progesterone was significantly elevated immediately after oviposition (stage 5), suggesting a possible role for progesterone in the regulation of sequential laying of egg pairs. J. Exp. Zool. 284:575-585, 1999.     

Domestication reduces the capacity to escape from predators

Phenotypic plasticity in response to variations in predatory pressure frequently occurs in wild populations, but it may be more evident and critical in species subjected to high exploitation rates and aquaculture. The Chilean scallop Argopecten purpuratus is becoming a domesticated species and the production of hatchery-reared scallops (closed environment), has implied the development of successive generations of individuals deprived of several stimuli normally present in their natural habitats (e.g. predators). We compared the escape capacities between wild and cultured A. purpuratus and also evaluated the effect of reproductive investment on the escape response capacities. Wild and cultured scallops, at different reproductive stages (maturing, mature and spawned), were stimulated to escape with the predatory sea star Meyenaster gelatinosus. We recorded: (1) the time to reaction, (2) the total number of claps, the duration of the clapping response and the clapping rate until exhaustion, (3) the time they spent closed after exhaustion, and (4) the proportion of claps recovered, the duration of the clapping response and the clapping rate after 20 min of recuperation. We found that wild A. purpuratus (1) reacted earlier when contacted by their natural predator, (2) escaped faster (greater clapping rates), (3) spent less time with their valves closed when exhausted, and (4) most of their escape capacities (i.e. claps number; clapping time; capacity of recuperation) were less affected by the energetic requirements imposed by gonad maturation and/or spawning than in cultured scallops. We considered that all these aspects of the escape response would make wild scallops less vulnerable to predation than cultured scallops, thus decreasing predation risk. Given the reduction of escape performance in cultured scallops, we suggest that this aspect should be considered for the success of culture-based restocking programs.     

The consequences of genetic variation in sex peptide expression levels for egg laying and retention in females

The accessory gland proteins (Acps) that male Drosophila melanogaster produce and transfer to females during copulation are key to male and female fitness. One Acp, the sex peptide (SP), is largely responsible for a dramatic increase in female egg laying and decrease in female receptivity after copulation. While genetic variation in male SP expression levels correlate with refractory period duration in females, it is unknown whether male SP expression influences female egg laying or if any effect of SP is mediated by SP retention in the female reproductive tract. Here we measured the amount of SP retained in the female reproductive tract after mating and female egg laying after copulating with virgin males. We found no correlation between male SP expression levels and egg laying, or the amount of SP in the female reproductive tract after mating. Additionally, the amount of SP retained in the female did not influence egg laying. These finding suggests that additional factors, such as variation in other Acps, are important for the retention of SP in females and its quantitative effects on egg laying. It also shows that egg laying and refractory period response to SP is at least partially uncoupled.     

Energy expenditure during egg laying is equal for early and late breeding free-living female great tits

In many bird populations, variation in the timing of reproduction exists but it is not obvious how this variation is maintained as timing has substantial fitness consequences. Daily energy expenditure (DEE) during the egg laying period increases with decreasing temperatures and thus perhaps only females that can produce eggs at low energetic cost will lay early in the season, at low temperatures. We tested whether late laying females have a higher daily energy expenditure during egg laying than early laying females in 43 great tits (Parus major), by comparing on the same day the DEE of early females late in their laying sequence with DEE of late females early in their egg laying sequence. We also validated the assumption that there are no within female differences in DEE within the egg laying sequence. We found a negative effect of temperature and a positive effect of female body mass on DEE but no evidence for differences in DEE between early and late laying females. However, costs incurred during egg laying may have carry-over effects later in the breeding cycle and if such carry-over effects differ for early and late laying females this could contribute to the maintenance of phenotypic variation in laying dates.     

The Female Post-Mating Response Requires Genes Expressed in the Secondary Cells of the Male Accessory Gland in Drosophila melanogaster

Seminal proteins from the Drosophila male accessory gland induce post-mating responses (PMR) in females. The PMR comprise behavioral and physiological changes that include increased egg laying, decreased receptivity to courting males, and changes in the storage and use of sperm. Many of these changes are induced by a “sex peptide” (SP) and are maintained by SP’s binding to, and slow release from, sperm. The accessory gland contains two secretory cell types with distinct morphological and developmental characteristics. Products of these “main” and “secondary” cells work interdependently to induce and maintain the PMR. To identify individual genes needed for the morphology and function of secondary cells, we studied iab-6^cocu males, whose secondary cells have abnormal morphology and fail to provide products to maintain the PMR. By RNA-seq, we identified 77 genes that are downregulated by a factor of >5× in iab-6^cocu males. By functional assays and microscopy, we tested 20 candidate genes and found that at least 9 are required for normal storage and release of SP in mated females. Knockdown of each of these 9 genes consequently leads to a reduction in egg laying and an increase in receptivity over time, confirming a role for the secondary cells in maintaining the long-term PMR. Interestingly, only 1 of the 9 genes, CG3349, encodes a previously reported seminal fluid protein (Sfp), suggesting that secondary cells may perform essential functions beyond the production and modification of known Sfps. At least 3 of the 9 genes also regulate the size and/or abundance of secondary cell vacuoles, suggesting that the vacuoles’ contents may be important for the machinery used to maintain the PMR.     

Fatty acid signatures of female Brünnich's guillemots Uria lomvia suggests reliance on local prey for replacement egg production

The laying of smaller replacement eggs has been described as a time-saving adaptation because chicks generally grow faster once hatched than inside the egg. However, chicks hatched from smaller eggs have, potentially, lower survival. Consequently, the smaller replacement egg represents a benefit to the female in terms of preserving its own condition at a cost to their offspring. We test these ideas by measuring adult mass changes and plasma lipid concentration changes in male and female Brünnich's guillemots Uria lomvia breeding on Coats Island, Nunavut. Though males lost more mass than females, these differences were not significant. Between laying the first and replacement egg, plasma fatty acid concentrations declined in females and increased in males, suggesting that females mobilise less lipid to preserve their condition after laying the replacement egg. In females, plasma lipid concentrations of the dominant fatty acids found in the eggs (16:0 and 18:1) declined between the laying of first and replacement eggs while plasma concentrations of 20:1 increased in both males and females. We compared the fatty acid signatures of first and replacement egg to look for evidence of differences between the lipid sources for their production. Principal component and discriminant function analyses showed that the fatty acid signatures of replacement eggs were closer to the signatures of the local prey than those of first eggs. We suggest that females rely on local sources of energy to a greater degree for the production of the replacement egg than the first egg, but that endogenous reserves of certain nutrients are important for the production of both eggs.     

Pigment-based skin colour in the blue-footed booby: an honest signal of current condition used by females to adjust reproductive investment

In monogamous species, the value of present reproduction is affected by the current condition of the mate, and females may use male ornaments to evaluate his condition and adjust their level of investment according. Many animals display colour in fleshy structures which may be accurate indicators of quality due to their potentially rapid response to changes in condition. Here we show that in the blue-footed booby, Sula nebouxii, male foot colour is structurally (collagen arrays) and pigment based. In 48 h foot colour became duller when males were food deprived and brighter when they were re-fed with fresh fish. Variation of dietary carotenoids induced comparable changes in cell-mediated immune function and foot colour, suggesting that carotenoid-pigmentation reveals the immunological state of individuals. These results suggest that pigment-based foot colour is a rapid honest signal of current condition. In a second experiment, we found that rapid variation in male foot colour caused parallel variation in female reproductive investment. One day after the first egg was laid we captured the males and modified the foot colour of experimental males with a non-toxic and water resistant duller blue intensive make-up, mimicking males in low condition. Females decreased the size of their second eggs, relative to the second egg of control females, when the feet of their mates were experimentally duller. Since brood reduction in this species is related to size differences between brood mates at hatching, by laying lighter second eggs females are facilitating brood reduction. Our data indicate that blue-footed booby females are continuously evaluating their mates and can perform rapid adjustments of reproductive investment by using dynamic sexual traits. We suggest that this fine-tuned adjustment may be widespread in socially monogamous animals.     

Sex-specific associations between reproductive output and hematozoan parasites of American kestrels

Parasites have the potential to decrease reproductive output of hosts by competing for nutrients or forcing hosts to invest in immune function. Conversely, reproductive output may affect parasite loads if hosts allocate resources to reproduction such that allocation to immune function is compromised. Both hypotheses implicitly have a temporal component, so we sampled parasites both before and after egg laying to examine the relationship between reproductive output (indexed using a combined measure of clutch size, egg volume, and initiation date) and blood parasite loads of American kestrels (Falco sparverius). Parasite loads measured prior to egg laying had no adverse effects on subsequent reproductive output. Females that previously had large reproductive outputs subsequently had lower parasite intensities than those whose outputs were smaller, suggesting that females were capable of allocating energy to both forming clutches and reducing parasite loads. Because male kestrels provide most of their mate's energetic needs before, during, and after egg laying, mate choice by females may have consequences for their parasite loads. Females choosing high-quality mates may not only have increased reproductive output, but may also obtain sufficient resources from their mates to enable them to reduce their parasite burdens. Males whose mates had large reproductive outputs were more likely to subsequently be parasitized and have more intense infections. For individual males sampled both before and after egg laying, those whose mates had larger reproductive outputs were also more likely to become parasitized, or remain parasitized, between sampling periods. Increased parasite loads of males may be one mechanism by which the costs of reproduction are paid.     

Interpopulation and context-related differences in responses of a marine gastropod to predation risk

We conducted laboratory experiments to investigate interpopulation differences in the behavioural responses of the whelk Buccinum undatum to the predatory lobster Homarus americanus and the asteroid Leptasterias polaris, both in the absence and presence of feeding opportunities. Whelks from three populations in the eastern North Atlantic (1) responded to lobsters by displaying avoidance behaviours (burrowing in the sediments or retreating inside their shell), (2) responded to asteroids by displaying escape responses (rapid crawling, shell rocking behaviour or foot contortions), and (3) more often refrained from feeding in the presence of a lobster than in the presence of an asteroid. Although whelks from the three populations responded similarly to lobsters and asteroids, interpopulation differences were evident. Thus, whelks from populations sympatric with a given predator more frequently displayed 'appropriate' antipredator behaviours (i.e. avoidance in the presence of a lobster, and escape in the presence of an asteroid) than did whelks allopatric with that predator. Also, whelks from a population sympatric with both predators fed less readily in the presence of a given predator than did whelks allopatric with that predator. However, the presence of a lobster or an asteroid had the same impact on the feeding response of whelks from two populations with contrasting predator fields, one sympatric with lobsters, but allopatric with asteroids, and one sympatric with asteroids, but allopatric with lobsters. The results of our study indicate that coexistence (over evolutionary or ecological time) with lobsters and asteroids increases the propensity of the whelk to display avoidance and escape behaviours in the presence of lobsters and asteroids, respectively, but has a less predictable effect on how whelks trade off predation risk and food acquisition. Studies are needed to investigate the roles of inheritance and experience on the development of antipredator behaviours and decision making by prey animals when predation risk conflicts with other fitness-related activities such as the acquisition of food or reproductive opportunities. Copyright 1999 The Association for the Study of Animal Behaviour.     

Life Cycle and Mating Behavior of Belonolaimus longicaudatus in Gnotobiotic Culture

The life cycle of Belonolaimus longicaudatus was observed in vitro on excised roots of Zea mays. Roots were cultured on Gamborg''s B5 medium in petri dishes with 1.5% agar adjusted to pH 5.8 and incubated at 28 °C in darkness. Second-stage juveniles (J2) fed on the roots and started the second molt (M2) to the third-stage juveniles 2 days after inoculation (DAI). The third molt (M3) to the fourth-stage juveniles occurred 7 DAI, followed by the fourth molt (M4) to males 13 DAI or to females 14 DAI. Nematode gender differences were observed by the end of the fourth molt. The first male appeared 15 DAI and the first female 17 DAI, after which mating occurred. Males were attracted to females, and mating was observed. Mating was required for reproduction. Fertilized females began to lay eggs 19 DAI and continued egg laying without the further presence of males during a 90-day observation. All of the eggs hatched. Unfertilized females rarely laid eggs, and none of the eggs were able to hatch. Feeding took place between each molt and before egg deposition occurred. The first-stage juveniles molted in the eggs 4 days after deposition, and J2 hatched from eggs 5 days after egg deposition. The life cycle from J2 to J2 was completed in 24 days.     

Biochemical content, energy composition and reproductive effort in the broadcasting sea star Asterias vulgaris over the spawning period

To assess sex differences in reproductive effort, we examined the biochemical composition and energetic content of the principal body components of the broadcast spawning sea star Asterias vulgaris in the Mingan Islands in the northern Gulf of St. Lawrence, eastern Canada. The body wall was the most stable body component, showing no variations in mass or in lipid and protein content (and total energetic content) between sexes or during spawning. Patterns in the gonads differed between sexes and with spawning. The lipid, protein and carbohydrate content of the ovary dropped during spawning, while only the protein content of the testis decreased significantly. Reproductive effort, expressed as loss of energy in the gonads during spawning for an individual weighing 10 g in underwater mass (8.2 cm in radius), was six times greater in females (49.5 kJ) than males (7.9 kJ). The energetic content of the pyloric caeca also decreased during spawning, by 17.7 kJ in females and 21.5 kJ in males, mainly due to a decrease in lipids. If this decrease is included as reproductive effort, it lessens the gender difference. The caecum decrease possibly represented expenditures due to formation of aggregations or the expulsion of gametes during spawning. Effectively, we observed aggregations during a massive spawning in this population. The sex ratio did not differ from 1:1 in all size classes sampled. This suggests that, unless males suffer higher mortality, females manage to allocate as much energy to somatic growth as males, possibly by feeding at higher rates to compensate for their higher reproductive effort. Stomach protein content tended to be higher in females than males and may indicate greater muscular development to facilitate digestion.     

Statural growth in known-age African elephants (Loxodonta africana)

The shoulder heights of 224 females and 170 males, and hindfoot length of 236 female and 217 male known-age African elephants ( Loxodonta africana ) were measured, and growth curves constructed for each measure of size. A linear relationship between foot length and shoulder height was confirmed in simultaneous measures of 97 males and 110 females. Growth curves demonstrated the typical sexual dimorphism in both foot length and shoulder height, with males growing more rapidly than females from birth onwards. The size dimorphism in foot length and shoulder height becomes marked by the age of 10 years, with males on average being 60–70 cm taller than females at 65 years. This size dimorphism is produced through faster growth which continues for longer than does that of females. The variance in growth rates is slightly greater for females than for males. It is proposed that female growth after puberty is affected by a trade-off between growth and reproduction, while males who deviate markedly from typical patterns of growth may be subject either to mortality or energetic constraints limiting their potential variance.     

Nocturnal and diurnal singing activity in the nightingale: correlations with mating status and breeding cycle

This study on the nightingale, Luscinia megarhynchos, is the first to examine both nocturnal and diurnal singing activity of mated and unmated males throughout a species' entire breeding cycle. Nocturnal song was sung mostly by unmated males. After pair formation, males ceased nocturnal singing and resumed it if their mate deserted. These results strongly suggest that nocturnal song of unmated males functions to attract a mate. Diurnal singing activity before females settled was low and did not predict future mating status. However, unmated males showed a continuous increase in diurnal singing activity until the end of the breeding cycle, but diurnal singing activity of mated males decreased after the egg-laying period. Mated males resumed nocturnal song for, on average, 3 nights during egg laying by their mates. This second period of nocturnal song coincided with the peak of diurnal singing activity. Such a high male singing effort during egg laying might allow the female to adjust her reproductive effort to male quality, deter rival males (e.g. through honest announcement of the female's fertility) or attract females for extrapair copulations. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Role of Catecholamines in the Reproduction of Romanomermis culicivorax

The relative concentrations of catecholamine in the nervous system of the entomophilic nematode Romanomermis culicivorax were measured under different experimental conditions by a glyoxylic acid-induced fluorescence procedure. A greater concentration of catecholamine was recorded in the nervous system of adult males and females than in postparasitic juveniles. A higher concentration of catecholamine occurred in adults maintained in physical contact with the opposite sex than in those maintained in isolation. Adult males maintained with females in the same aqueous medium but physically separated by a barrier displayed a greater concentration of catecholamine in their nervous systems than did males maintained in isolation, but the catecholamine fluorescence intensity of such males was less than in males allowed physical contact with females. In adult males, the fluorescence intensity of catecholamine declined progressively during and after copulation. In adult females, the intensity of catecholamine remained constant before, during, and after copulation. Catecholamine(s) may play a role in regulating copulatory behavior, egg production, or oviposition.     

Squid egg mops provide sensory cues for increased agonistic behaviour between male squid

Males of many species compete for access to females. In order to avoid performing potentially costly agonistic behaviour for their entire adult lives, many group-living males use environmental cues to limit agonistic behaviour to times when it will be of most benefit. Long-finned squid, Loligo pealeii, live less than a year and aggregate in mixed- and single-sex schools. Adults participate in several spawning events, then die. During spawning events, males actively compete for females. Winning males pair with females, which subsequently lay eggs in communal sites on the ocean floor (‘egg mops’). To determine whether males use sensory cues provided by egg mops to regulate agonistic behaviour, we conducted four laboratory experiments. We measured the agonistic responses of pairs of adult males before, during and after exposure to conspecific egg mops. In three experiments, egg mops were manipulated to provide differing sensory stimuli (tactile, water-borne, visual). The addition of conspecific egg mops to tanks of paired male squid dramatically increased agonistic behaviour above control levels within minutes. Male squid were first attracted to the egg mops visually, but contact with the capsules was necessary to increase agonistic behaviour. After initial contact, agonistic behaviour was almost continuous as long as egg mops remained present, even when squid touched the egg mops infrequently. Visual stimuli seemed important in maintaining elevated agonistic behaviour between egg mop touches. When egg mops were removed from the tank, measured agonistic behaviour declined within minutes. When egg mops were added to the tank while covered by an opaque and porous cover that allowed water-borne stimuli to circulate into the tank, squid did not approach the covered egg mop or show increased agonistic behaviour. This result suggests that water-borne stimuli are not sufficient to increase agonistic behaviour. It is unusual for male agonistic behaviour in any species to be increased by contact with fertilized eggs. In this species, however, egg capsules might signal that sexually mature, receptive females are about to lay eggs. Indirect evidence suggests that mating with a female immediately before she lays eggs increases male paternity. If this prediction is true, the presence of egg mops may indicate the optimal time for male squid to establish mating precedence through agonistic bouts. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Egg size and reproductive adaptations among Arctic deep-sea copepods (Calanoida,<Emphasis Type="Italic"> Paraeuchaeta</Emphasis>)

Reproductive strategies of the four congeneric and sympatric calanoid copepods Paraeuchaeta glacialis, P. norvegica, P. barbata, and P. polaris were studied in the Arctic Greenland Sea. Females of all species produce egg sacs and carry their brood attached to the genital opening until the offspring hatch. However, egg size and lipid content as well as clutch size and the fraction of females carrying egg masses show characteristic differences among the four species. P. glacialis and P. norvegica produce large numbers (37 to more than 50) of relatively small eggs, whereas P. barbata and P. polaris rely on small numbers (10 to 19 and 4 to 6, respectively) of large eggs with a high energy content. There is no correlation between female body size and egg size or clutch size, respectively. Females of the smallest species, P. polaris, produce relatively large eggs and show the highest energetic investment per egg. In contrast, energetic investment per clutch is highest in P. glacialis. Reproductive adaptations appear to be strongly related to the depth range inhabited by the respective species. In the central Greenland Sea P. glacialis and P. norvegica occur in the epipelagial and upper mesopelagial, whereas the other two species inhabit lower mesopelagic to bathypelagic depths. Thus, egg size increases with depth of occurrence, whereas clutch size is inversely correlated with depth. This observation leads to the hypothesis that the evolution of large eggs in deep-sea copepods may enable hatchlings to rely on a lecithotrophic development and thus represents a successful adaptation to cope with the limited food supply at great depths, whereas high offspring numbers in epipelagic species compensate for higher predation risks in the euphotic zone.Communicated by H.-D. Franke