Estimate of removal rate ofNereis virens (Polychaeta: Nereidae) from an intertidal mudflat by gulls (Larus spp.)

The feeding behavior of herring gulls (Larus argentatus), ringed-billed gulls (L. delawarensis) and great blackbacked gulls (L. marinus) on an intertidal mudflat in Maine, USA, was investigated. Remains of fish, mussels, crabs, insects, and the polychaeteNereis virens were recovered from gull feces. Forty-three percent of the fecal samples containedN. virens jaws, setae, or both. A comparison of jaws from fecal material and from worms collected from the natural community demonstrates that gulls preferentially preyed upon larger worms. Feeding was largely confined to 3 h around low tide, and birds fed mostly in the low intertidal and below mean low water where the largest worms were located. Individual birds remained on the flat for a mean of 28.0 min and consumed a mean of 19.2 worms per visit. It was calculated that gulls remove a mean of 808 largeN. virens from the flat per tide from June to October, representing an estimated 0.04% of the standing crop of largeN. virens.     

Histochemical demonstration of mercury in the intestine,nephridia and epidermis of the marine polychaete Nereis virens exposed to inorganic mercury

Marine polychaetes Nereis virens (Sars) were exposed to 9 ppb Hg as mercuric chloride in the aquarium water. Concentrations of mercury in the water and in the worms were monitored using radiolabelled mercury. After 11 d, the average mercury concentration in the nereids was 8.41 ppm Hg and the bioconcentration factor was calculated to be 930. Accumulations of mercury in the tissues were made visible for light and electron microscopy by autometallographic silver enhancement. Mercury was demonstrated in the intestine, nephridia, epidermis and cuticula. In the intestine, mercury was found to be located predominantly in the apical part of the epithelial cells. In the nephridia, mercury deposits were apparent in the peritoneal cells of the nephridial tubules. Silver-enhanced mercury was also observed at the epicuticula, in the collagen fibres of the endocuticula and in the epidermal basal cells. Intracellularly, the observed mercury accumulations were localised predominantly in lysosomes. Extracellularly, mercury was observed in the basal laminae of the intestine and the epidermis, and also in the intestinal peritrophic membrane. How N. virens copes with mercury toxicity is discussed.     

Social influence on the correlation between behaviours in young-of-the-year perch

The connection between risk-taking behaviour and exploratory behaviour in young-of-the-year perch (Perca fluviatilis) was studied in aquarium experiments to see whether individual behaviour patterns could be identified in this species and also to investigate how individual behaviour is influenced by their social environment. Risk-taking was defined as the time spent foraging in an open area vs hiding in the vegetation in the presence of a piscivore. Explorative behaviour was measured as latency to enter a passage leading to an unknown area. Groups of four fish were used for the observations, and both behaviours measured were positively correlated with the mean scores of these behaviours in the other group members. Risk-taking and explorative behaviours were correlated only when data was adjusted for the behaviour of the other group members. Individuals that spent more time in the open than their companions also tended to be faster than the others to enter the passage to the unknown area and vice versa. The results indicate that there are consistent individual differences in boldness in perch, but also that behaviour could be modified according to the behaviour of group members.     

Migration of the sandworm Nereis virens during winter nights

There have been many previous reports of the sandworm Nereis virens Sars swimming in the water column. This behavior usually has been attributed to reproductive processes. Sandworms were found swimming in surface waters at night on ebb tides during many nights of January, February and March in a Maine (USA) estuary. None of the specimens examined contained gametes or possessed other characteristic spawning or pre-spawning modifications. Several age classes were found, with worms measuring 9 to 38 cm in length, weighing 0.5 to 19.8 g, and having 82 to 187 segments. The greatest numbers of worms were observed during near-average tides on evenings in which low tides occurred a few hours after sunset but prior to moonrise. Up to 83 worms per minute were observed swimming seaward through a 20 m transect, while none were observed swimming landward at any stage of the tide. It is concluded that sandworms swimming during winter nights is unrelated to reproduction and that it is an inherent behavior pattern.Contribution No. 102 of the Ira C. Darling Center, Walpole, Maine 04573, USA.     

Microbial reaction rates and bacterial communities in sediment surrounding burrows of two nereidid polychaetes (Nereis diversicolor and N. virens)

The effects of infaunal mode of life on sediment properties, microbial reaction rates, as well as abundance and composition of bacterial communities were studied in sediment surrounding burrows (mucus lining, oxidised wall, ambient anoxic and surface sediment) of two closely related, but behaviourally different, nereidid polychaete worms: the facultative suspension-feeder Nereis (Hediste) diversicolor and the obligate deposit-feeder Nereis (Neanthes) virens. Burrow sediment of the two species was collected from two adjacent (50 m distance) shallow sandy locations (Kertinge Nor, Denmark). The burrow lining and wall of both polychaete species were enriched in organic matter originating from mucous secretions by the inhabitants and phytoplankton trapped through irrigation. This was more evident for N. diversicolor that shows a significantly higher irrigation rate than N. virens. Both the organic matter mineralisation rates (based on anaerobic incubations) and bacterial abundance were higher along the burrow linings and walls. However, accumulation of porewater TCO₂ and dissolved organic carbon in sediments adjacent to burrows increased most rapidly in the presence of N. diversicolor, suggesting higher heterotrophic activity associated with this species. Surprisingly, bacterial abundance was lower around burrows of N. diversicolor than those from N. virens indicating that burrow environments from the first species harbour a more active bacterial community. Molecular fingerprints of the 16S rRNA gene from bacterial communities showed that the composition of the burrow linings and walls resembled the ambient anoxic sediment rather than the oxic sediment surface. On the other hand, the bacterial fingerprints of the sediment surrounding the burrows of the two polychaete species were markedly different suggesting either a site-specific difference in sediment parameters or a significant species-specific impact of the burrow inhabitants.     

Pollutant effects on the mixed function oxygenase (MFO) and reproductive systems of the marine polychaeteNereis virens

The cytochrome P-450 dependent mixed-function oxygenase (MFO) system ofNereis virens, found in the endoplasmic reticulum of the intestinal epithelium, has similar components and cofactor requirements when compared to other invertebrates. Worms exposed to food containing benzo(a)pyrene or a mixture of polychlorinated biphenyls (Aroclor 1254) had significantly higher MFO activity and cytochrome P-450 content than unexposed ones.N. virens from a site in Porland, Maine, USA, which was heavily contaminated with oil, and nearby clean reference sites were studied. The weight, MFO activity and presence or absence of gametes were determined. The MFO activity in worms from the oil-contaminated site was approximately six times that of worms from the reference sites; worms from the reference areas were six times larger than oil-contaminated worms. Worms from the oil-contaminated area lacked or had undeveloped gametes in spring, summer and fall. In contrast, many of the worms collected from the reference sites at the same times had fully developed gametes.     

Foraging behaviour of sympatric Antarctic and subantarctic fur seals: does their contrasting duration of lactation make a difference?

The duration of periods spent ashore versus foraging at sea, diving behaviour, and diet of lactating female Antarctic (Arctocephalus gazella, AFS) and subantarctic (A. tropicalis, SFS) fur seals were compared at Iles Crozet, where both species coexist. The large disparity in lactation duration (SFS: 10 months, AFS: 4 months), even under local sympatry, has led to the expectation that AFS should exhibit higher foraging effort or efficiency per unit time than SFS to allow them to wean their pups in a shorter period of time. Previous evidence, however, has not supported these expectations. In this study, the distribution of foraging trip durations revealed two types of trips: overnight (OFT, <1 day) and long (LFT, >1 day), in common with other results from Macquarie Island. However, diving behaviour differed significantly between foraging trip types, with greater diving effort in OFTs than in LFTs, and diving behaviour differed between fur seal species. OFTs were more frequent in SFS (48%) than in AFS (28%). SFS performed longer LFTs and maternal attendances than AFS, but spent a smaller proportion of their foraging cycle at sea (66.2 vs. 77.5%, respectively). SFS dove deeper and for longer periods than AFS, in both OFTs and LFTs, although indices of diving effort were similar between species. Diel variation in diving behaviour was lower among SFS, which foraged at greater depths during most of the night time available than AFS. The diving behaviour of AFS suggests they followed the nychthemeral migration of their prey more closely. Concomitant with the differences in diving behaviour, AFS and SFS fed on the same prey species, but in different proportions of three myctophid fish (Gymnoscopelus fraseri, G. piabilis, and G. nicholsi) that represented most of their diet. The estimated size of the most important fish consumed did not vary significantly between fur seal species, suggesting that the difference in dive depth was mostly a result of changes in the relative abundance of these myctophids. The energy content of these fish at Iles Crozet may thus influence the amount and quality of milk delivered to pups of each fur seal species. These results contrast with those found at other sites where both species coexist, and revealed a scale of variation in foraging behaviour which did not affect their effort while at sea, but that may be a major determinant of foraging efficiency and, consequently, maternal investment.     

Photoperiodic control of growth and segment proliferation by Nereis (Neanthes) virens in relation to state of maturity and season

The effects of two fixed photoperiodic regimes on the rate of somatic growth and segment proliferation in the semelparous polychaete Nereis (Neanthes) virens Sars has been investigated. The two photoperiods (16 h light:8 h dark and 8 h light:16 h dark, hereafter LD 16:8 and LD 8:16) were close to the extremes experienced naturally by N. (N.) virens in the course of a year. The experiments were carried out during the ambient winter and all worms (including four-setiger larvae raised out of season) showed elevated feeding, growth and enhanced segment-proliferation rates when reared under LD 16:8, compared to worms reared under LD 8:16. The rate of replacement of lost segments after caudal ablation was also higher under LD 16:8. The number of segments present when the segment proliferation rate had fallen to zero was higher in individuals grown from birth under LD 16:8 compared to those grown under LD 8:16. Feeding activity showed a degree of spontaneous recovery under LD 8:16 from January onwards, but remained lower than under LD 16:8. This difference continued into the following spring, and was not attributable to differences in sexual maturity. We conclude that the rate of feeding in N. (N.) virens and other indicators of somatic growth rate such as rates of caudal regeneration and segment proliferation are directly influenced by the photoperiod, but that the seasonal cycle of growth also includes a circa-annual component modifying the response to static photoperiods according to the time of year. Received: 20 October 1998 / Accepted: 4 February 1999     

Hydrolytic enzymes in the coelomic cells of the polychaeteNereis virens during sexual maturation

Nereis virens were collected between April 1989 and April 1990 at Yerseke, Oosterscheldt Bay, The Netherlands. Activities of the hydrolytic enzymes arylsulfatase, acid phosphatase, N-acetylglucosaminidase, aminopeptidase, and carboxylesterase were investigated in coelomic cells (elaeocytes) of individuals at different stages of maturation as determined by the time course of oocyte growth. On a protein-content basis, up to ten-fold higher specific activities were present in the elaeocytes compared to the body-wall tissue. Acid phosphatase, N-acetylglucosaminidase, leucine aminopeptidase, and long- and medium-chain esterase increased continuously from the beginning to the later stage of sexual maturation, with a decline in prespawning individuals. Shortchain esterase and acid aminopeptidase did not show corresponding changes. Activities at different maturation stages were similar for males and females, with the exception of arylsulfatase activity, which was sex-specific and almost lacking in male elaeocytes. The high hydrolytic activities present in the elaeocytes as well as the increase in these activities during sexual maturation are in agreement with the function of these cells in phagocytizing muscle fragments (sarcolytes), which arise as a result of drastic histolysis of the body musculature ofN. virens during the process of sexual maturation.     

Sex differences in the foraging behavior of squirrel monkeys in a seasonal habitat

Summary The effects of sex and seasonal changes in food abundance on foraging behavior was studied in squirrel monkeys (Saimiri oerstedi) in Costa Rica over an eleven-month period. Females searched for and ate food at significantly greater frequencies than did males throughout the study. The frequency of the specific foraging techniques used occasionally differed significantly within seasons, but not across the study period. Few differences were found in the foraging behaviors of nonreproductive sexually mature females compared to females that were pregnant or lactating. The major exception was that during the month following parturition reproductive females foraged for flowers and fruits more frequently than did non-reproductive females. The reduction of time spent by males in foraging activities gives them more time for other activities, especially anti-predator vigilance. Foraging techniques and the proportions of different food types in the diet changed seasonally. Foraging for arthropods was most frequent in the season when arthropod abundance was lowest, resulting in the amount of time spent eating arthropods to vary less across the seasons. Fruits and flowers were not eaten in a direct relationship to availability, but were used more than expected relative to availability when arthropod abundance was reduced. Individuals were more dispersed when foraging compared to other activities. Overall, there was little evidence of any direct foraging benefits for a squirrel monkey from being social.     

Modified pheromone communication associated with insecticidal resistance in the obliquebanded leafroller, Choristoneura rosaceana (Lepidoptera: Tortricidae)

Summary. This study reports on the impact of insecticidal resistance on the diel periodicity of the calling behaviour and pheromone production of different-aged virgin females of the obliquebanded leafroller (OBL), Choristoneura rosaceana. While both resistant (R) and susceptible (S) females initiated calling on the first night following emergence, the periodicity of the calling behaviour, as determined by the mean onset time of calling (MOTC) and the mean time spent calling (MTSC) over the first six nights of calling, differed between the two strains. R females started calling significantly later in the night. However, as the MOTC of R females advanced with age but did not do so in S individuals, the difference between strains was more pronounced in younger than older females. Furthermore, R females spent less time calling than S individuals. However, the MTSC increased as a function of age in both R and S females, so the difference between strains remained fairly constant for each night of calling. The major component of OBL sex pheromone, the Z11-14: Ac, determined at peak calling activity, significantly declined with female age. Overall, pheromone production was lower in R females than in S females, with the difference being more pronounced in younger than in older individuals. Thus, resistant females may have a lower mating success. The mating success of both R and S strain males did not vary with the number of previous matings acquired. With regard to males, although there was a significant decline in spermatophore size with successive matings, there was no significant difference between strains. However, R males are smaller and may be disadvantaged through female choice and/or may respond differently to pheromone source compared with S individuals. If the reproductive success of both sexes is affected, this may have a profound influence on the dynamics of insecticidal resistance in the presence or absence of selection in OBL populations. Received 4 July 2001; accepted 19 October 2001.     

Different tactics,one goal: initial reproductive investments of males and females in a small Arctic seabird

Despite a great number of studies on extra-pair paternity in birds, the actual roles of males and females in extra-pair contacts is poorly understood, as detailed behavioural studies comparing the reproductive performance of the two sexes prior to egg laying are relatively scarce. Here, we investigated mating behaviour (copulations and aggressive interactions), time budget and body condition (size-adjusted body mass and baseline corticosterone level) in the little auk (Alle alle), a monogamous and highly colonial, Arctic seabird. We performed the study in a large breeding colony of the little auk in Hornsund (Spitsbergen). We found that the males frequently attempted extra-pair copulations (EPCs), although these contacts were almost always unsuccessful, mostly because of the females’ rejection behaviour. These results clearly indicate that genetic monogamy is maintained through female control. Nevertheless, males tried to protect their paternity by staying in close proximity to their females and aggressively intervening when their mates became involved in EPCs. Compared to females, males also spent more time in the colony guarding nest sites. Despite the apparent sex differences in the time budget and frequency of aggressive interactions, body condition was similar in the two sexes, indicating comparable parental investments during the mating period.     

Bilaterality in the regular sea urchin Anthocidaris crassispina is related to efficient defense not to efficient locomotion

Regular sea urchins show pentaradial symmetry: they have round bodies made of five almost identical sectors. There is no preference in which sector should be anterior in their locomotion. Anthocidaris crassispina is exceptional among regular sea urchins. Morphometry showed that their tests had radial symmetry but the overall body shape showed bilateral symmetry because of the uneven distribution of spine length. This bilateral axis did not correspond to the ambulacrum III—interambulacrum 5 axis, which is the bilateral axis of irregular sea urchins. Their behavior in relation to this bilaterality was observed in the aquarium. At rest, most of them stayed in the upper corner of the aquarium keeping the sector with the shortest spine length (the shortest sector) toward the water surface. At walking, most of them moved on the vertical wall just under the water surface keeping the shortest sector directing to the water surface. When they walked in the open space, they moved in a random direction with respect to the bilaterality of the spines. The bilaterality and the behavior of this sea urchin were interpreted as the adaptation to defense not to efficient locomotion.     

Effect of temperature on activity and social behavior of the adult tautog Tautoga onitis under laboratory conditions

Experiments were conducted on the effects of high, sublethal temperature on activity, social behavior, and feeding in the adult tautog Tautoga onitis held under controlled laboratory conditions. Norms of these behaviors were established during acclimation to temperatures at which these fish are commonly found. During acclimation, the fish were active and feeding throughout the day. A dominance hierarchy, based primarily on size, was established for each group, with subordinate fish being displaced from all areas of the aquarium. Under elevated temperature, changes in established norms of these behaviors were observed, with activity, feeding, aggression, and the spacing between fish showing a decrease. Following return to pre-rise temperatures, although aggression remained reduced, activity, interfish distance, and feeding approached levels established during acclimation. A comparison of these results with previous ones on young tautog is made and the significance of the findings is discussed.This work was supported, in part, by a grant from the U.S. Energy Research and Development Administration No. E(49-7) 3045.     

Importance of predation by infaunal polychaetes in controlling the structure of a soft-bottom community in Maine,USA

Field manipulations in a Maine, USA intertidal mud flat showed that Nereis virens adults were an important factor in regulating the abundance of Corophium volutator, an amphipod which comprised 63% of the total number of individuals in the benthic community. Removal of N. virens adults resulted in an increase in C. volutator abundance, while addition of N. virens adults led to a decrease. There was a significant negative correlation between the abundance of N. virens adults and that of C. volutator. C. volutator was also negatively correlated with the density of all the other infaunal species combined. These results suggest that there is at least a 3-level interactive system (N. virens —C. volutator — other infaunal species) within the infauna in Maine; N. virens is a factor in controlling the abundance of C. volutator, and C. volutator may play a role in regulating the densities of other infaunal species. Models of generalized cropping in soft-bottom systems may be too simple if they ignore complex trophic relationships within the infauna.     

Reactions of fish to sound generated by divers' open-circuit underwater breathing apparatus

Observations were made on the influence of underwater divers on the behaviour of fish in Loch Torridon, on the west coast of Scotland. An electronically scanned sonar and a television camera were used to record the behaviour of 4 species common in the loch. These were the cod Gadus morhua (L), the saithe Pollachius virens (L), the pollack P. pollachius (L) and the common dab Limanda limanda (L). These fish were attracted to divers and to a sound source when the recorded noise from an aqualung and demand valve was transmitted into the water. By analysis and by testing different components of the noise it was shown that low frequencies between 30 and 110 Hz, generated by the release of exhaled air, were responsible for the attraction. It is suggested that the fish associated this noise with the presence of food organisms disturbed from the sea bed by the diver and that they had become conditioned to the noise over a period of time.     

Bilateral symmetry and locomotion: do elliptical regular sea urchins proceed along their longer body axis?

Vagile animals usually have bilaterally symmetrical bodies and proceed with their mouth-end first. Regular sea urchins have, however, radially symmetrical bodies with their mouth facing the substratum and show no preference in which side of the body should be anterior in their locomotion. The elliptical sea urchins in the subfamily Echinometrinae are exceptional among regular sea urchins in having elongated bilateral bodies. We studied whether they showed the preference in the direction of locomotion using Echinometra sp. type A. Directional preference was not observed in the proceedings in an open space. However, they proceeded preferentially with their long axis coinciding with the direction of locomotion when they moved along the water surface or along the wall of the aquarium. The speed of locomotion was the same irrespective of the direction of proceedings and of whether sea urchins moved freely or moved along the water surface or along the walls. We suggest that the bilateral body form and the habit of long-axis lead of this sea urchin have adaptive significance to increase the protected body surfaces, not to facilitate the efficiency in locomotion.     

Satellite tagging of blue sharks (Prionace glauca) and other pelagic sharks off eastern Australia: depth behaviour, temperature experience and movements

Satellite telemetry was used to study the movements and behaviour of ten blue sharks and one individual each of shortfin mako, thresher and bigeye thresher off eastern Australia. The tracks showed latitudinal movements of up to 1,900 km, but none of the sharks travelled away from the eastern Australian region. Tracking periods did not exceed 177 days. All species showed oscillatory dive behaviour between the surface layers to as deep as 560–1,000 m. Blue sharks spent 35–58% of their time in <50 m depths and 10–16% of their time in >300 m. Of these four species, the bigeye thresher spent the least time in the surface layers and the most time at >300 m depth. All four species showed clear diel behaviour generally occupying shallower depths at night than during the day. Blue sharks were mainly in 17.5–20.0°C water, while the thresher sharks showed a more bimodal temperature distribution.     

Locating food in a spatially heterogeneous environment: Implications for fitness of the macrodecomposer Armadillidium vulgare (Isopoda: Oniscidea)

To assess the fitness consequences of foraging on patchy resources, consumption rates, growth rates and survivorship of Armadillidium vulgare were monitored while feeding in arenas in which the spatial distribution of patches of high quality food (powdered dicotyledonous leaf litter) was varied within a matrix of low quality food (powdered grass leaf litter). Predictions from behavioural experiments that these fitness correlates would be lower when high quality food is more heterogeneously distributed in space were tested but not supported either by laboratory or field experiments. To investigate whether A. vulgare can develop the ability to relocate high quality food patches, changes in foraging behaviour, over a comparable time period to that used in the fitness experiments, were monitored in arenas in which there was a high quality food patch in a low quality matrix. A. vulgare increased its ability to relocate the position of high quality food over time. It reduced time spent in low quality food matrices and increased time spent in high quality food patches with time after the start of the experiment. When the position of a high quality food patch was moved, the time spent in the low quality food matrix increased and less time was spent in high quality food patches, compared to arenas in which the food was not moved. The fitness benefits for saprophages of developing the ability to relocate high quality patches while foraging in spatially heterogeneous environments are discussed.     

Experimental evidence of a testosterone-induced shift from paternal to mating behaviour in a facultatively polygynous songbird

Previous studies have suggested that testosterone (T) profiles of male birds reflect a trade-off between mate attraction behaviours (requiring high T levels) and parental care activities (requiring low T levels). In this study, we experimentally elevated T levels of monogamous males in the facultatively polygynous European starling (Sturnus vulgaris), and compared mate attraction and paternal behaviour of T-treated males with those of controls (C-males). T-males significantly reduced their participation in incubation and fed nestlings significantly less often than C-males. Females paired to T-treated males did not compensate for their mate’s lower paternal effort. The observed reduction in a male’s investment in incubating the eggs was accompanied by an increased investment in typical female-attracting behaviours: T-males spent a significantly higher proportion of their time singing to attract additional females. They also occupied more additional nestboxes than C-males, although the differences just failed to be significant, and carried significantly more green nesting materials into an additional nestbox (a behaviour previously shown to serve a courtship function). T-males also behaved significantly more aggressively than C-males. During the nestling period, the frequency of mate-attracting behaviours by T-treated and control males no longer differed significantly. Despite the reduced paternal effort by T-males and the lack of compensation behaviour by females, hatching and breeding success did not differ significantly between T- and C-pairs. Received: 7 February 2000 / Revised: 10 August 2000 / Accepted: 3 September 2000