Interactions between sea urchins (Strongylocentrotus droebachiensis) and their predators in field and laboratory experiments

Field observations and manipulative experiments in a nearshore cobble bed (2 to 3 m below mean low water) at Eagle Head, Nova Scotia, Canada, between 1984 and 1986, showed that small juveniles ofStrongylocentrotus droebachiensis (3 to 6 mm diam) sheltering beneath cobbles had a refuge from predators such as rock crabs, small lobsters, and fish. Sea urchins gradually outgrew these refuges and small adults (25 to 30 mm) required larger rocks as shelter from predators, particularly large cancrid crabs. Small juveniles were usually solitary and well dispersed beneath cobbles, whereas small adults tended to aggregate on the undersides and in the interstices of boulders. These aggregations may develop passively as sea urchins accumulate in suitablysized refuges. Chemotaxis experiments indicate that juvenileS. droebachiensis are repelled by waterborne stimuli from conspecifics. In a factorial experiment, effects of the presence of potential predators (rock crabs and lobsters) and/or food (kelp) on the behaviour of large juvenile (10 to 15 mm) and small adult sea urchins were examined in flowing seawater tanks. Both size classes formed exposed feeding aggregations when kelp was provided as food, irrespective of the presence or absence of predators. In the absence of kelp, each size class responded differently to the presence of a predator: juveniles became more cryptic, whereas adults aggregated on the tank sides. Increased movement to the sides of a tank in the presence of a predator may reflect a flight response, since chemotaxis experiments indicated thatS. droebachiensis is repelled by waterborne chemical stimuli from predators. Observational and experimental data suggest that predation at the late juvenile and early adult stages may influence population structure, distribution and abundance ofS. droebachiensis.     

Vocalizations reveal body condition and are associated with visual display traits in great frigatebirds (Fregata minor)

Acoustic displays are known to advertise aspects of male quality and to affect female choice of mates in a variety of birds, including not only songbirds but some seabirds as well. Male great frigatebirds (Fregata minor) produce a rapid warble vocalization that forms a prominent part of their courtship display. We investigated the relationships between aspects of this vocalization and male quality and pairing success in 103 great frigatebirds from a population in the northwestern Hawaiian Islands. We found that across males, frequency bandwidth of the display was negatively related to repetition rate, enabling us to use deviation from the upper bound regression of bandwidth on rate (??vocal deviation??) as one measure of vocal performance. We used vocal consistency, defined as the coefficient of variation of the time interval between successive peaks in the warble, as a second measure of performance. Vocal deviation was significantly positively associated with body condition of male great frigatebirds, but vocal consistency was not. We found that peak frequency of the warble was not significantly associated with body size but was negatively associated with size of the gular pouch, the most prominent visual ornament. Male great frigatebirds showed significant individual differences in all three vocal measures??peak frequency, vocal deviation, and vocal consistency??but none of these measures was a significant predictor of pairing success in our study population. These results suggest that vocalizations provide honest information about male body condition and gular pouch size in great frigatebirds but do not influence male success in pairing with females.     

Effect of intraspecific density on life history traits and population growth rate of Neanthes arenaceodentata (Polychaeta: Nereidae) in the laboratory

The effects of intraspecific density on life history traits and population dynamics of the nereid polychaete Neanthes arenaceodentata Moore were assessed in a laboratory experiment. Survival, growth, and fecundity were measured for one generation of worms at densities of 40, 80 and 160 worms per 840 cm² (1x, 2x, and 4x treatments, respectively). Density did not affect size (prior to pairing), percentage of worms paired, time to pairing, or size of mature paired males. Density did have a significant negative effect on survival, size of mature paired females, time to spawning, percentage of females that reproduced, and number of eggs per reproducing female. As density increased, mean survival was 90.0, 80.8 and 74.0%; mean size of mature females was 52.2, 49.2 and 48.1 segments; mean time to spawning was 100.6, 102.4 and 109.4 d; and mean fecundity was 881, 622, and 598 eggs per female, for 1x, 2x and 4x treatments, respectively. Increased density reduced the potential population growth rate, ; for a given rate of larval survivorship, was lower in the 2x and 4x treatments than the 1x treatment. Analysis of sensitivity of to changes in survivorship indicated that population growth rate at the highest density was sensitive to both changes in larval survivorship and the probability of producing a successful brood, although at low densities, was sensitive only to changes in larval survivorship. We attribute these density effects to aggressive bahavioral interactions between the worms, primarily the adults. This experiment identifies key life history traits that could be measured in future experiments to test population level responses of N. arenaceodentata populations to pollutants, both in the laboratory and in the field.Contribution No. 820 of the U.S. Environmental Protection Agency, Environmental Research Laboratory and Contribution No. 189 of the Marine Sciences Institute, University of Connecticut     

Sexual signalling and viability in a wolf spider (Hygrolycosa rubrofasciata): measurements under laboratory and field conditions

This study examined the crucial prediction of the conditional-handicap theory, the relationship between male sexual trait size and male viability, in the wolf spider Hygrolycosa rubrofasciata. In this species, males court females by drumming dry leaves with their abdomen, and males with the highest drumming rate enjoy highest mating success. We determined male drumming rate, body mass, and mobility, which reflects mate-searching activity, in relation to male survival. Because it is often difficult to know how results obtained from laboratory studies reflect the natural world, particularly when the measured variable is survival, we repeated our study in both laboratory and field conditions. Males drumming at the highest rate survived better than males drumming at a lower rate in both laboratory and field conditions. These results are in accordance with the predictions of conditional-handicap models of sexual selection. Survival was independent of male body mass and there was no significant correlation between male drumming activity and body mass. However, large males moved further than smaller males, and males moving longer distances lost less mass than those moving shorter distances. These results suggest that, moving, and consequently mate-searching activity, may be a condition-dependent trait and that there may be a advantage for large males in mate searching. Received: 22 October 1998 / Received in revised form: 28 January 1999 / Accepted: 14 February 1999     

Genetic protein variation in red mullet (Mullus barbatus) and striped red mullet (M. surmuletus) populations from the Mediterranean Sea

Starch-gel electrophoresis of allozymes was used to differentiate the two red mullet species (Mullus barbatus L. and M. surmuletus L.) in the Mediterranean Sea and, further, to investigate the genetic stock structure of M. barbatus in the eastern Mediterranean area. Twenty putative enzyme-coding loci were examined in eight M. barbatus samples caught in the Aegean and Ionian Seas (Greece) and in the Gulf of Lion (France), and two M. surmuletus samples caught in the Aegean and Gulf of Lion. A high degree of genetic polymorphism was found in both species. Species-specific electrophoretic patterns were found in PGI* and PGM*. Estimates of variance of allele frequencies among samples (F _ST) and ² analyses both revealed significant differences (P < 0.05) among the M. barbatus samples. Most of the genetic variation was among samples regardless of region. The mean value of Nei's genetic distance between the two species was 0.329. Genetic distance among M. barbatus samples was low (maximum Nei's D = 0.012), with the sample from Platania differing most from other M. barbatus samples. This is probably be due to founder effects existing at this area. These results suggest that allozyme analysis may provide important information on the genetic structure of the red mullet to ensure sustainable management of this species. Received: 7 May 1997 / Accepted: 13 October 1997     

Mating behaviour of the tiger blue butterfly (Tarucus theophrastus): competitive mate-searching when not all females are captured

Summary Females of the tiger blue butterfly (Tarucus theophrastus) fly upwind in search of the larval hostplant. Males perch or patrol the downwind edges, searching for incoming females or those already on the bush. A model of competitive mate-searching is developed for the case where not all receptive females are contacted by searching males: the model differs from the earlier ones of Parker (1970, 1974), particularly when few males are involved. Observed distributions of males upon bushes of different sizes agree better with the predictions of the game theory model than with a random distribution of males. The described model may be widely applicable to populations where females remate, but frequently evade detection by searching males.     

Estimating paternity with spatial behaviour and DNA fingerprinting in the striped plateau lizard, Sceloporus virgatus (Phrynosomatidae)

The association between spatial proximity and paternity was studied in a population of the striped plateau lizard, Sceloporus virgatus. The relationship between estimated mating success and male phenotypic traits was examined for a sample of 55 males. DNA samples were obtained from 13 female-offspring families. The males with the closest spatial proximity to each female were tested as possible sires within each family. Fingerprinting with two multilocus hypervariable minisatellite probes revealed a strong correspondence between male-female spatial proximity and actual paternity. Paternity could be assigned for 72 of the 100 hatchlings. Most hatchlings with identifiable sires were attributed to a male with the highest category of spatial proximity to the mother. However, there was a low to moderate level of multiple paternity within clutches, and for some clutches probable sires could not be identified even though the most likely behavioural candidates were tested. Thus, nonterritorial males or other males lacking strong social and spatial relationships with females may achieve some degree of reproductive success. Analysis of mating success revealed that male success increased with body size, up to a point beyond which larger size conferred no advantage. Received: 7 January 1997 / Accepted after revision: 16 June 1997     

Dietary selectivity in the field and food preferences in the laboratory for two herbivorous fishes (Cebidichthys violaceus and Xiphister mucosus) from a temperate intertidal zone

Diets and food selectivity of two stichaeid fishes (Cebidichthys violaceus and Xiphister mucosus) from the rocky intertidal zone of the central California coast (USA) were studied at each season of the year by gut content analysis and abundance measurements of potential macrophyte food items. Both fishes, after reaching a standard length of about 44 mm, were almost exclusively herbivorous. The bulk of the diet consisted of 8 to 10 species of chlorophytes and rhodophytes. These main dietary components were chiefly annual seaweeds with high surfaceto-weight ratios (sheetlike forms or small, highly branched forms). Perennial seaweeds were eaten in relatively large amounts only during the winter. Macrophytes eaten in only trace amounts included about 20 species of chlorophytes, chrysophytes, phaeophytes, rhodophytes and a spermatophyte. The small amount of animal material in the diet (never more than 2% by weight) could well have been ingested incidentally while the fishes were feeding on seaweeds. Food preference tests with up to 19 macrophyte species in the laboratory revealed that both fishes chose to eat three annual rhodophytes (Smithora naiadum, Porphyra perforata and Microcladia coulteri) in preference to Ulva lobata, an annual chlorophyte that was more abundant in the diets of field-caught specimens.     

Growth and reproduction ofCaprella danilevskii (Crustacea: Amphipoda) reared in the laboratory

Caprella danilevskii Czerniavski, an epifaunal amphipod crustacean, was successfully reared for two generations under laboratory conditions. Amphipods were maintained in pairs or fours in a Petri dish. Temperature and photoperiod were maintained at 20°C, and 14 h light: 10 h dark, respectively. Colonies of the diatomCylindrotheca closterium (Ehrenberg) Reumann and Lewin were provided as food source. The red algaGelidium amansii (Lamouroux) Lamouroux was used to provide an attachment substrate for the caprellids. Juveniles that emerged from the brood pouch were described as instar I. Subsequent instars were identified by molts. Juveniles molted successively at 2.5 to 6.5 d intervals. The body length of males at each instar increased exponentially, while that of females followed a sigmoid curve. A different pattern of flagellar segment addition in the antenna I of males and females was revealed. At the age of 20.8 d, females reached instar VII, and produced their first embryos. After 5.1 d, juveniles emerged from the brood pouch. On average, females produced 5.4 clutches successively at 5.0 d intervals. The number of offspring per spawning increased from 6.3 at instar VII to 22.0 at instar XII. The total number of offspring was 69.0 for a female throughout her life span.     

Variation in condition of coastal Newfoundland 0-group Atlantic cod (Gadus morhua): field and laboratory studies using simple condition indices

Demersal 0-group Atlantic cod (Gadus morhua L.) were collected in Trinity Bay, Newfoundland, Canada, between August 1993 and January 1994. Diet composition and simple indices of condition (hepato-somatic index, condition factors and body ash content) were determined. Laboratory experiments were also conducted to determine whether simple indices of condition reflect differences in energy intake (ration level) and to determine minimum index of condition values, assessed from 0-group cod that died due to exhaustion of their energy reserves. Determination of lower critical indices of condition provided a meaningful interpretation of field data on feeding, condition and survival of 0-group cod. In the wild, indices of condition reflecting variation in liver and muscle energy reserves of cod increased rapidly with the consumption of Calanus finmarchicus, which exhibited a high incidence of oil sacs, and declined abruptly when this lipid-rich prey was no longer consumed. Liver and muscle condition of 0-group cod from the field varied seasonally, but their condition was consistently above that of cod that died in the laboratory. Laboratory studies, using three ration levels, revealed that condition indices differed over time and among ration levels. Indices of condition representing variation in muscle energy reserves exhibited a steady decline once the hepato-somatic index fell below about 2.0. We interpret this as evidence for depletion of lipids first and then proteins, a pattern similar to that of larger juveniles and adults. We conclude that as long as 0-group cod are above a “safe” level of condition their body length can still increase even when condition is declining. Received: 2 September 1997 / Accepted: 8 December 1998     

Near real time satellite tracking of striped marlin (Kajikia audax) movements in the Pacific Ocean

High-resolution satellite locations were obtained from striped marlin using Argos transmitters attached to the upper lobe of the caudal fin. Twenty-six striped marlin were tagged off New Zealand (2005–2007) and tracked as far as the central Pacific Ocean. Caudal fin mounted Argos tags generated 1,524 locations during a total of 659 tracking days [mean 25 (±21.24) days per fish and 2.3 (±2.30) locations per day]. 38% of locations have an estimated accuracy of ±1 km or better. Displacement rates from high quality locations ranged from 2.9 to 170.8 km in a 24 h period, with a mode at 20–30 km and a mean of 45 km/day. The caudal fin attachment methodology and antenna configuration was adjusted each season to improve transmission life and data quality, with the best results obtained in the last year of deployments (2007). The longest track duration was 102 days, with a total displacement of 4,959 km and a total track distance from all locations received of 6,850 km. Tag shedding and antenna failure appear to have limited the duration of tracks from SPOT tags. The high temporal and spatial resolution data revealed behaviours not previously observed in striped marlin, including associations to subsurface bathymetric features. High resolution location data such as these are useful inputs for statistical models used to investigate habitat selection and switching between different behavioural modes. The geolocations calculated using ukfsst estimates from PAT tag data had RMS errors of 1.01° latitude and 0.59° longitude when compared with SPOT tag Argos locations.     

Personality trait differences between mainland and island populations in the common frog (Rana temporaria)

Understanding and predicting species range expansions is an important challenge in modern ecology because of rapidly changing environments. Recent studies have revealed that consistent within-species variation in behavior (i.e., animal personality) can be imperative for dispersal success, a key process in range expansion. Here we investigate how habitat isolation can mediate differentiation of personality traits between recently founded island populations and the main population. We performed laboratory studies of boldness and exploration across life stages (tadpoles and froglets) using four isolated island populations and four mainland populations of the common frog (Rana temporaria). Both tadpoles and froglets from isolated populations were bolder and more exploratory than conspecifics from the mainland. Although the pattern can be influenced by possible differences in predation pressure, we suggest that this behavioral differentiation might be the result of a disperser-dependent founder effect brought on by an isolation-driven environmental filtering of animal personalities. These findings can have important implications for both species persistence in the face of climate change (i.e., range expansions) and ecological invasions as well as for explaining rapid speciation in isolated patches.     

Maturation,epitoky and regeneration in the polychaete Eunice siciliensis under field and laboratory conditions

In the gonochoric polychaetous annelid Eunice siciliensis Grube, germ cells only develop in the posterior segments of the worm; these segments subsequently differentiate into an epitokous portion separating at maturity from the infertile body part. As in the Pacific and Atlantic palolo worms, the epitokous portions die some hours after spawning. Laboratory investigations revealed that the anterior parts of spawned worms survive and regenerate genital segments. After a period of at least 8 months, paired gonads are developed in these segments. Gonia proliferating later from the germinal tissue are released from gonadal sacs into the coelomal cavities, where gamete development occurs. A reproductive epitokous portion does not mature and separate until the second year of cultivation; it is followed by caudal regeneration. These facts and the observation that only about 50% of the individuals obtained from samples of coralligène were sexually differentiated, led to the conclusion that E. siciliensis probably has a bi-annual reproductive cycle. Caudal regeneration does not depend on the presence of a prostomia hormone. The ability to form posterior regenerates is inherent to the parapodial segments of all body regions, except for mature genital segments. This is also valid for the formation of morphologically different secondary prostomia. However, for initiation and further promotion of gametogenesis and epitokous development, at least one gonadotropic factor originating from the prostomium is required. Neurosecretory cells within the brain suggest this factor(s) to be of neuro-endocrine nature. Specimens developing secondary prostomia were found both under field and laboratory conditions. In several cases, sexual development took place, and even epitokous portions were separated from such individuals. From this it appears that the gonadotropic functions are re-established by the regenerated prostomia in order to ensure reproduction in individuals accidentally deprived of the anterior end. The possible rôle of temperature and light as environmental factors involved in the control of normal and regenerative growth, of sexual development and reproduction is discussed.     

Photosynthetic oxygen production and PAM fluorescence in the brown algaPadina pavonica measured in the field under solar radiation

The effects of solar radiation on photosynthetic oxygen production, pulse amplitude-modulated (PAM) fluorescence and pigmentation were measured in the Mediterranean brown macroalgaPadina pavonica (Linnaeus) Lamouroux under field conditions and natural sunlight. Exposure of thalli to solar radiation for 1 h caused a dramatic decrease of their photosynthetic quantum yield, which recovered to initial levels after they had been placed in the shade for 3 h. Photoinhibition also occurred at the natural growth site ofP. pavonica during the hours of maximal solar irradiance. Photosynthetic oxygen production was also affected by high levels of solar radiation both in algae harvested from the surface and from 6 m depth; oxygen production started to decrease after a few minutes of exposure, and negative values were found after 1 h of solar exposure. Chlorophylla content inP. pavonica also decreased during the hours of maximal solar irradiation. These results suggest that photoinhibition ofP. pavonica occurs during part of a typical summer day on Mediterranean coasts.     

Natural selection and the communal rearing of pups in African wild dogs (Lycaon pictus)

Summary The contributions of adult (>18 months) and yearling (10–18 months) African wild dogs (Lycaon pictus), to raising pups were quantified at eight natal dens, where pups remain for their first three months of life.Adults between 2–6 years of age did most of the hunting, and the dominant male often made the first grab at fleeing prey. Yearlings contributed to the hunting but were reluctant to tackle large prey animals. Yearlings and breeding females had prior access to food. Yearlings and adults regurgitated comparable amounts of food, but in one pack watched at a time of food shortage, the yearlings failed to regurgitate and stole food from the pups.Dominant dogs chased predators, especially spotted hyenas (Crocuta crocuta) from the area of the den more frequently than other adults or yearlings chased predators. Mothers, particularly in the first six weeks post-partum, stayed and protected the pups when the pack hunted.There is a positive, but non-significant correlation between the number of adult helpers and the number of pups raised. However, the sex ratio bias towards males at birth suggests that male helpers (which predominate) usually increase pup survivorship. The roles of direct and indirect selection in the evolution of the helping behavior are discussed. Indirect selection has probably played an important role in the unusual post-reproductive survival of males and their helping behavior.Serengeti Wildlife Research Institute contribution number 284