Differences in stress tolerance and brood size between a non-indigenous and an indigenous gammarid in the northern Baltic Sea

Differences in stress tolerance and reproductive traits may drive the competitive hierarchy between non-indigenous and indigenous species and turn the former ones into successful invaders. In the northern Baltic Sea, the non-indigenous Gammarus tigrinus is a recent invader of littoral ecosystems and now occupies comparable ecological niches as the indigenous G. zaddachi. In laboratory experiments on specimens collected between June and August 2009 around Tvärminne in southern Finland (59°50′N/23°15′E), the tolerances towards heat stress and hypoxia were determined for the two species using lethal time, LT₅₀, as response variable. The brood size of the two species was also studied and some observations were made on maturation of juveniles. Gammarus tigrinus was more resistant to hypoxia and survived at higher temperatures than G. zaddachi. Brood size was also greater in G. tigrinus than in G. zaddachi and G. tigrinus matured at a smaller size and earlier than G. zaddachi. Hence, there are clear competitive advantages for the non-indigenous G. tigrinus compared to the indigenous G. zaddachi, and these may be further strengthened through ongoing environmental changes related to increased eutrophication and a warming climate in the Baltic Sea region.     

Reproductive patterns of three intertidal salt-marsh gammaridean amphipods

Among three sympatric species of epibenthic amphipods found at different tide marks at Jamaica Bay, New York (USA), the length of time juveniles spend with the mother increases and the number of juveniles per brood decreases as tidal height increases. Each brood has two developmental periods: (1) the embryonic period, from ovulation to hatching; (2) the juvenile period, from hatching to emergence from the marsupium. Gammarus palustris, found at the high-tide mark, has a mean juvenile period of 1.7 days and a mean brood size of 12.4 offspring; G. mucronatus, found at mean-tide mark, has a mean juvenile period of 0.8 days and a mean brood size of 27.4 offspring; Melita nitida, found at low-tide mark, has a mean juvenile period of 0.5 days and a mean brood size of 30.0 offspring. Further, the range of days that a juvenile may emerge is widest for G. palustris (0 to 8 days after hatching) and narrowest for M. nitida (0 to 2 days).This work was submitted in partial fulfillment of the requirements for the Ph.D. at the City University of New York.     

Light responses of the larva of the serpulid polychaeteGaleolaria caespitosa

AdultGaleolaria caespitosa were collected at Cape Banks and Balmoral Bay, both in the vicinity of Sydney, Australia, from October 1985 to February 1986. Planktotrophic larvae of this serpulid polychaete are negatively phototactic at irradiance levels between 10 and 2 000x10¹⁴ q cm^-2 s^-1. The effective spectral range lies below 530 nm. Early metatrochophore larvae are less strongly photonegative; later they become photopositive. The photopositive phase persists into the actively swimming 3-setiger stage. Older setigerous larvae are photoneutral, demersal and positively thigmotactic. The strength of the photonegative response is not affected by the level of irradiance. The photopositive response is weaker at low levels of irradiance. The photonegative behaviour of young larvae may keep them at an optimal level for feeding. The subsequent photopositive phase may increase chances of being washed ashore and the terminal, photoneutral, thigmotactic phase may be important in the selection of a suitable site for settlement. Comparisons are made with light responses of two other serpulid species. Larval responses to light may be important infitness, especially in relation to successful niche occupation by adults.     

In vitro reproduction of the turbellarian<Emphasis Type="Italic"> Urastoma cyprinae</Emphasis> isolated from<Emphasis Type="Italic"> Mytilus galloprovincialis</Emphasis>

Urastoma cyprinae is a species infecting the gills of several marine bivalves. Although there is some literature on this turbellarian, its life cycle remains unknown. In our work we have demonstrated that reproduction of U. cyprinae can be completed out of its host. More than 50% of turbellarians isolated from mussels (Mytilus galloprovincialis) secreted and cemented a cocoon to the well bottom during the first 72 h of incubation in seawater (34 salinity) at 14°C. Oviposition started at days 1–18 (average 4.8 days) and occurred in most cases inside the protective cocoon. Each Urastoma laid an average of 2.9 egg capsules (range 1–10) and 3.9 embryos were developed inside each egg capsule (range 1–11). Hatching started at days 20–43 (average 24 days). An average of 12.8 juvenile forms (range 1–64) escaped from the cocoon after hatching. The free-swimming juveniles showed a positive phototactic response and survived about a month after hatching. On the basis of our results, we propose a life cycle for U. cyprinae involving a sexual maturation parasitic period in the bivalve gills and a reproduction period including cocoon secretion, egg laying, and hatching that is entirely completed in the external environment.Communicated by S.A. Poulet, Roscoff     

Phototaxis in Fungiidae corals (Scleractinia)

Corals within the scleractinian family Fungiidae were observed to move toward light (positive phototaxis). Negative phototactic movement was not observed in any of the specimens tested. On sandy substratum, Diaseris distorta moved faster (max. speed of 3 cm h^-1) than other species. Although D. distorta has symbiotic algae, phototactic movement also was observed both in bleached corals and in those treated with a specific inhibitor (dichlorophenyl dimethyl urea) of photosynthesis. D. distorta was phototactic even on a glass plate, and climbed up a steep slope (up to 30°). Based on experiments with Fungia fungites and D. distorta, soft tissues at the peripheral region of the dise seem to be responsible for movement via peristalsis. It is suggested that positive phototaxis in symbiontbearing fungiid corals is an important trait for selection of favorable habitats.     

Comparative study of crustacean larval photoresponses

Ovigerous females of four brachyuran (Cancer gracilis, Lophopanopeus bellus bellus, Hemigrapsus oregonensis and Scyra acutiforns) and two anomuran (Pagurus beringanus and P. granosimanus) species, which live as adults in coastal areas, were collected near Friday Harbor, Washington, USA, in 1985, and spectral sensitivity and phototactic pattern of their larvae were measured. Responses were compared with previous measurements on estuarine species to determine whether responsiveness varies with adult habitat. Estuarine and coastal species have similar photoresponses. Spectral sensitivity of the test brachyran species had two maxima, one near 400 to 420 nm and another around 500 nm. The anomuran species were similar, but had an additional peak in the region of 580 to 620 nm. This sensitivity is adapted to daytime light conditions in the adult environment, and available spectra at the time of larval movement during diel vertical migration. Upon light adaptation and stimulation with a narrow light field, all six species showed positive phototaxis to high light intensities and a pronounced negative response to low intensities. Only the negative response will occur in natural underwater light conditions, and it is part of a predator-avoidance shadow-response which operates in areas of higher light intensity. The same phototactic pattern is observed in all species when darkadapted except H. oregonensis. If nocturnal vertical migration occurs, this negative response may be responsible for the descent at sunrise and depth maintenance during the day.     

Transfer of symbiotic luminous bacteria from parental Leiognathus nuchalis to their offspring

Juveniles of Leiognathus nuchalis were raised from fertilized eggs for up to 60 d and examined for luminescence activity. Almost all juveniles raised separately from adults failed to produce detectable light. In contrast, a significant percentage (33 to 100%) of the juveniles became luminescent in less than 48 h when they were either kept with adults or inoculated with a homogenate of the adult light organs. The luminescence tended to increase with time after the treatments. These findings suggest that: (1) most of L. nuchalis offspring typically hatch and develop apo-symbiotically and (2) at least 45 d after hatching, juveniles can be infected with symbiotic luminous bacteria from the light organ of adult fish, and thereby gain the ability to produce light. Received: 16 February 1999 / Accepted: 21 September 1999     

Ontogeny of space use and diet of two temperate damselfish species, <Emphasis Type="Italic">Parma microlepis</Emphasis> and <Emphasis Type="Italic">Parma unifasciata</Emphasis>

Studies of reef fish herbivory have mainly focused on the impacts and behaviour of adults of tropical species. In this study, the ontogenetic shifts in home range, aggression, feeding rate, diet and gut morphology in juveniles and adults of two temperate territorial damselfishes, Parma microlepis and Parma unifasciata, were determined. Both P. microlepis and P. unifasciata juveniles under 80 mm TL exhibited no aggressive chases towards conspecifics or other species, while above 80 mm TL aggressive chase frequency increased in conjunction with an increase in home range, defended as a territory. Ontogenetic diet shifts, characterised by an increase in herbivory (P. unifasciata: juveniles: 64% plant material, adults: 95% plant material; P. microlepis: juveniles: 43% plant material, adults: 67% plant material) were observed for both species. The ratio of digestive tract length to body length, which often accompanies a switch to herbivory, increased significantly with ontogeny for both species. Compared to tropical confamilial grazers, these temperate damselfish species feeding rates were lower, and they had larger territories which were not as strongly defended (fewer aggressive chases).     

Patterns of co-occurrence and interactions between age classes of the common triplefin,Forsterygion lapillum

Many marine organisms have pelagic larval stages that settle into benthic habitats occupied by older individuals; however, a mechanistic understanding of intercohort interactions remains elusive for most species. Patterns of spatial covariation in the densities of juvenile and adult age classes of a small temperate reef fish, the common triplefin (Forsterygion lapillum), were evaluated during the recruitment season (Feb–Mar, 2011) in Wellington, New Zealand (41°17′S, 174°46′E). The relationship between juvenile and adult density among sites was best approximated by a dome-shaped curve, with a negative correlation between densities of juveniles and adults at higher adult densities. The curve shape was temporally variable, but was unaffected by settlement habitat type (algal species). A laboratory experiment using a “multiple-predator effects” design tested the hypothesis that increased settler mortality in the presence of adults (via enhanced predation risk or cannibalism) contributed to the observed negative relationship between juveniles and adults. Settler mortality did not differ between controls and treatments that contained either one (p = 0.08) or two (p = 0.09) adults. However, post hoc analyses revealed a significant positive correlation between the mean length of juveniles used in experimental trials and survival of juveniles in these treatments, suggesting that smaller juveniles may be vulnerable to cannibalism. There was no evidence for risk enhancement or predator interference when adults were present alongside a heterospecific predator (F. varium). These results highlight the complex nature of intercohort relationships in shaping recruitment patterns and add to the growing body of literature recognizing the importance of age class interactions.     

Locomotion,feeding, grooming and the behavioural responses to gravity,light and hydrostatic pressure in the stage I zoea larvae of Ebalia tuberosa (Crustacea: Decapoda: Leucosiidae)

The stage I zoeae of Ebalia tuberosa swam by sculling with the exopodites of the 1st and 2nd maxillipeds and flexed the abdomen to brake or change direction. The larvae gained depth by stopping all natatory movements and sinking passively at rates of 6 mm s^-1. The zoeae refused both living and dead nauplii of Artemia spp., as well as two species of diatoms, but fed readily on detritic material on the bottom which they scooped up using the endopodites of the maxillipeds and pressed against the mouthparts using the telson. The setae on the posterior border of the telson were used for grooming the maxillipeds and the anterior mouthparts. Day-old stage I zoeae were negatively geotactic, positively phototactic and responded to pressure increases by swimming upwards and by high barokinesis. By the third day some larvae had become positively geotactic but were photopositive, and the majority responded to pressure increases as in the day-old larvae. Five-day old larvae were still photopositive but the majority had become positively geotactic and fewer himbers responded to pressure. Seven-day old larvae failed to respond to any of the stimuli used and assumed a predominantly benthic lifestyle. It is suggested that this anomalous behaviour is related to the dispersal of the larvae and to the specialized habitat requirements of the adults while the rather unusual morphology of the larvae is related to their feeding behaviour and semi-benthic lifestyle.     

Larval behavior of predacious sister-species: orientation,molting site,and survival in Chrysopa

Field and laboratory studies compared two features of larval behavior in a pair of predacious sisterspecies of green lacewings: one (Chrysopa slossonae) a specialist on a single species of colonial aphids (the woolly alder aphid) that occur on branches and trunks of alder trees, the other (C. quadripunctata) a general aphid feeder whose primary prey is dispersed on foliage of diverse types of trees. First, a few hours after hatching, larvae of the two species develop significantly different phototactic responses; the differences correspond well with the spatial distributions of their prey. Most C. slossonae exhibited negative phototaxis, a response that helps move hatchlings inward on alder trees toward the woolly alder aphid colonies, whereas most C. quadripunctata hatchlings showed positive orientation to light, a response that tends to keep them in tree canopies with their prey. Second, in greenhouse experiments, a significantly greater proportion of C. slossonae larvae (second instars) molted within woolly alder aphid colonies and remained with the aphids than did C. quadripunctata larvae. These differences indicate that the specialist larvae have evolved a high degree of behavioral fidelity to their prey. However, larvae (second instars) of the two species that were released near ant-tended woolly alder aphid colonies in the field had similar recovery (= survival) rates. Consequently, natural selection may not act on behavioral traits that influence larval fidelity to prey during the late second and early third instars.     

Sex,cannibalism and sticklebacks

Summary Two closely related species of sticklebacks (Pisces:Gasterosteidae), breeding in the same site at the same time, showed different patterns of egg cannibalism. In Gasterosteus aculeatus the intensity of cannibalism was density-dependent and females were the most important cannibals. In G. wheatlandi cannibalism was not density-dependent and males ate more eggs than females. This study indicates that sex-specific patterns of cannibalism can differ widely between species despite their similar ecology, and it provides evidence inconsistent with a currently popular explanation for egg cannibalism.     

Fitness consequences of sexual cannibalism in female<Emphasis Type="Italic"> Argiope bruennichi</Emphasis>

The evolution of sexual cannibalism as the most extreme form of nuptial feeding is still poorly understood. Although increasing evidence suggests that female aggressiveness is related to other aspects of foraging behaviour, it is not clear whether the nutritional value of a male is sufficient to provide an adaptive significance for sexual cannibalism. A widely cited though rarely tested explanation is based on a paternal investment model, and predicts that consumption of a male results in increased female fecundity. The available evidence is either correlational or restricted to species with relatively large and potentially nutritious males, and different studies have come to different conclusions. Here we present a test of the paternal investment hypothesis using the very cannibalistic and highly size-dimorphic spider Argiope bruennichi. After a preset schedule, we had females consume none, one or two males independent of the female's cannibalistic behaviour. Consumption of male bodies did not result in any detectable fitness benefit for the female: neither the number of clutches, nor clutch size or hatching success were affected by consumption of males. The frequency of cannibalism was around 80%, independent of the female mating status. We did not observe male complicity, but cannibalism was associated with prolonged copulation. This suggests a sexually selected benefit of cannibalism for males. We conclude that the paternal investment hypothesis does not explain the existence of sexual cannibalism in A. bruennichi and probably not in other spider species with a pronounced sexual size dimorphism.Communicated by L. Simmons     

Precopulatory sexual cannibalism in fishing spiders (Dolomedes triton): a role for behavioral syndromes

Precopulatory sexual cannibalism (predation of a potential mate prior to copulation) offers an extreme example of intersexual conflict, a current focus in behavioral ecology. The aggressive-spillover hypothesis, posits that precopulatory sexual cannibalism may be a nonadaptive by-product of a general syndrome of voracity (aggression towards prey) that is expressed in multiple behavioral contexts. In this view, selection favoring high levels of voracity throughout ontogeny spills over to cause sexual cannibalism in adult females even when it is not necessarily beneficial. Using the North American fishing spider, Dolomedes triton, we present the first in depth test of this hypothesis. We found support for three aspects of the spillover hypothesis. First, voracity towards hetero-specific prey results in high feeding rates, large adult size, and increased fecundity. Second, juvenile and adult voracity are positively correlated (i.e., voracity is a consistent trait over ontogeny). Third, voracity towards hetero-specific prey is indeed positively correlated with precopulatory sexual cannibalism. Assays of antipredator behavior further revealed positive correlations between boldness towards predators, voracity and precopulatory sexual cannibalism. Overall, our results support the notion that precopulatory sexual cannibalism in D. triton is part of a behavioral syndrome spanning at least three major contexts: foraging, predator avoidance, and mating.     

Tisbe pori n. sp. (Copepoda: Harpacticoida) from the Mediterranean coast of Israel and its cultivation in the laboratory

Tisbe pori n. sp., from the Mediterranean coast of Mikhmoret, Israel, was cultivated in the laboratory and is described. The relative food values of 9 species of marine algae were evaluated according to the copepods' development time (length of period from hatching to reproductive maturity). Individuals fed on Dunaliella tertiolecta or D. primolecta developed the fastest. The rate of feeding of the adults on Dunaliella primolacta was determined. In the temperature range 17° to 31.5°C, the shortest development time (7 days) was at 26° to 28°C. The range of salinity tolerated by adults was 26 to 48.     

Respiration of Sepia officinalis during embryonic and early juvenile life

Adult Sepia officinalis L. were caught in June 1984, in the coastal waters of Wimereux (France). Deposition of the eggs took place in the seawater aquaria of the Station Marine. The oxygen consumption of S. officinalis was measured during embryonic and juvenile development. Aerobic metabolism occurs as soon as the early embryonic Stage 21. Oxygen diffuses through the initially thick egg shell; the oxygen level in the perivitelline liquid reaches a maximal value just before hatching (116.7±6.9 mm Hg). Hatchings display only a slight increase in oxygen consumption compared to embryos in the last stage of development. Respiration experiments with 40 d old juveniles showed that oxygen consumption increases with temperature, but is not affected by photoperiod. Experiments under increasing hypoxia revealed that S. officinalis juveniles are good regulators and maintain a constant oxygen consumption in the range of 4 to 7 mg O₂l^-1. Juveniles successfully recover from an hypoxic stress of 2 mg O₂l^-1 maintained for 1 h. This suggests that the respiratory pigments (pre-hemocyanins) of 40 d-old juveniles have a high oxygen affinity and/or that these juveniles have the ability to adapt to anaerobic conditions.     

Larval development in the Antarctic nemertean Parborlasia corrugatus (Heteronemertea: Lineidae)

Embryonic and larval development were followed from fertilisation to settlement in the Antarctic heteronemertean Parborlasia corrugatus (McIntosh, 1876). The first cleavage occurred 10 to 15 h after fertilisation, and the second at 17 h. Larvae hatched at the gastrula stage, between 170 and 200 h post-fertilisation, and were 150 m in diameter. Early larval stages aggregated in dense groups near the surface of incubation vessels and were positively phototactic. Early pilidium larvae were recognisable 435 h post-fertilisation. They were 155×152 m in size, and possessed a complete apical tuft of cilia and a full marginal band of locomotory cilia. At this stage, the gust was visible through the body wall, and the mouth was open and was 40 m in diameter. Late pilidia, 222×193 m in size, were helmet-shaped. They had an apical tuft over 100 m long, and possessed a lobed marginal band of locomotory cilia. Pilidia were observed aggregating close to the bottom of incubation vessels 1200 to 1350 h (50 to 56 d) after fertilisation, and this was interpreted as settlement behaviour. At this stage, the apical tuft had been lost and they were highly contractile, being capable of compressing their bodies. However, neither developing juveniles within the larval envelope nor hatched juveniles were observed. Pilidia consumed the microalgae Tetraselmis suecica, Thalassiosira pseudonana and Isochrysis galbana. They also fed on particulate organic material < 1 m in size, as shown by the presence of material in the guts of larvae offered filtered extracts of algal cultures. There was some indication that larvae could use dissolved organic material, since pilidia held in seawater with organic material removed did not survive as long as those in filtered seawater or in filtered water with added amino acids. However, the only larvae to exhibit settlement behaviour in the feeding experiments were those offered Tetraselmis succica and Thalassiosira pseudonana, and these required a longer development time to reach this stage than pilidia in the standard cultures, where a mixed algal diet was offered.     

A test on social behavior as a cause of dispersal of spruce grouse

Summary Social behavior is one hypothesized cause for dispersal. We tested this hypothesis on spruce grouse (Dendragapus canadensis) in two years by removing adults in summer and monitoring emigration of juveniles in autumn. Despite the absence of adults, juveniles emigrated from the natal range at rates comparable to juveniles on control areas. Data on density and dispersal from earlier spruce grouse studies are reviewed, which show that emigration is constant despite substantial differences in density, and that a clear female sex bias occurs across a wide range of densities. Results do not support the hypothesis that social behavior influences dispersal nor do they support a prediction of the Oedipus hypothesis that no sex bias occurs in polygynous birds. Offprint requests to: D.M. Keppie     

Length-weight relationships and coefficient of condition of Euphausia superba and Thysanoessa macrura (Crustacea: Euphausiacea) in southwest Indian Ocean during summer

Samples of antarctic euphausiids, Euphausia superba and Thysanoessa macrura were obtained during the MD 25 FIBEX expedition of the R. V. Marion Dufresne, in February 1981, to the southwest Indian Ocean. Individual and mean coefficients of condition (K _m) were calculated from variations in weight as a function of total length and of carapace length. In E. superba, no significant differences were found as a function of sex or development stage in either total length and weight or carapace length and weight. In T. macrura, a significant allometric difference emerged in the wet weight of juveniles, males and females as a function of total length, and in wet weight as a function of carapace length between adults and subadults. In both E. superba and T. macrura, wet weight as a function of total length differed significantly among stations, and also between species, with a greater weight increase for T. macrura in summer. The coefficients of condition calculated for all wet weight-total length relationships showed that the morphologically different Group II males were heavier than Group I males and mature females. K _m in T. macrura was higher for females than for juveniles or males, indicating a greater weight gain by the females. These differences probably reflect real differences in physiology and may affect the distribution of these two species.     

Comparative sensitivity of three life stages of the tropical mysid,Metamysidopsis insularis to six toxicants

Test protocols citing mysids as a standard test species often recommend juveniles (48 h) as the most appropriate life stage for testing. Acute toxicity tests were conducted with six standard toxicants, with three life stages of Metamysidopsis insularis; juveniles (48 h), late juveniles (7–12 days) and adults (>13 days), to determine whether there were significant differences in the sensitivity between life stages. Three of the compounds tested (sodium dodecyl sulphate, potassium dichromate, and potassium chloride) showed a significantly higher 96 h-LC₅₀ for the adults when compared to the juveniles and late juveniles. Four of the compounds tested showed significantly higher 24 h- and 48h-LC₅₀ for the adults when compared to both the late juveniles and juveniles. Twenty-four hour-LC₅₀ values for juveniles were also significantly higher than the 48 and 96 h values. However, for the late juveniles and adults this was only true of a few compounds. Further, the 48 and 96 h values were statistically similar. Therefore, for M. insularis it may be appropriate to conduct acute toxicity test with juvenile for a 48 h period.