Divergence in female calling song discrimination between sympatric and allopatric populations of the southern wood cricket Gryllus fultoni (Orthoptera: Gryllidae)

Two species of closely related wood cricket, Gryllus fultoni (Orthoptera: Gryllidae) and Gryllus vernalis, occur together in some parts of the eastern United States and have a similar calling song structure, consisting of three-pulse chirps. A previous study revealed that chirp rate in G. fultoni was highest (greatest difference vis-à-vis chirp rate in G. vernalis) in sympatric populations, intermediate in near allopatric populations that were located close to the sympatric zone, and lowest in allopatric populations. A similar trend was observed in pulse rate, but the mean values of this trait showed much more convergence than chirp rate at the low end of the range of calling temperatures. In this study, we investigated the song discrimination of females from sympatric and allopatric populations of G. fultoni at about 23°C, which is near the middle of the normal range of calling temperatures. We used both single-stimulus and two-stimulus playback experiments to learn if geographical differences in song preferences paralleled those in calling songs. Stimuli presented were representative of calling songs in three classes of G. fultoni populations (sympatric, near allopatric, and far allopatric), a calling song of G. vernalis, and three calling songs with parameter values that were intermediate with respect to those of the songs of far allopatric G. fultoni and G. vernalis. In the single-stimulus playbacks, females of all G. fultoni populations responded poorly if at all to the heterospecific stimulus. Females of sympatric and near allopatric populations responded poorly to all intermediate stimuli, but females of far allopatric populations frequently responded to these sounds. In the two-stimulus playbacks, females of sympatric and near allopatric populations generally discriminated against intermediate and heterospecific stimuli. However, females of far allopatric populations often did not discriminate against intermediate stimuli, whose characteristics resembled the calling songs of G. vernalis. The divergent pattern of female phonotactic discrimination between sympatric and far allopatric populations was thus generally congruent with the pattern of divergence in chirp and pulse rates and would be expected to significantly reduce heterospecific mating in sympatry. These geographical patterns of female song discrimination and male calling songs conform to a commonly used definition of reproductive character displacement.     

Assessment and individual recognition of opponents in the pygmy swordtails Xiphophorus nigrensis and X. multilineatus

We examined factors that determine the outcome of agonistic encounters between male pygmy swordtail fish. Xiphophorus nigrensis and X. multilineatus males formed dominance relationships based on body size in staged laboratory encounters. There was a significant negative correlation between size asymmetry and fight intensity, suggesting that males assessed size in the encounters. However, a significant proportion of the variation in fight intensity in contests that escalated to bites could not be explained by size asymmetry. Aggressive motivation may also influence the outcome of contests and could be assessed in agonistic encounters. Theory suggests that signals of aggressive intention will be evolutionarily stable if individuals can recognize opponents and encounter one another repeatedly. In addition, individual recognition is one way that dominance hierarchies can be maintained. Here we demonstrate that males from both species can recognize individuals. In addition, at least some X. nigrensis males were site-faithful in the field, suggesting males encounter the same opponents repeatedly.Communicated by G.M. Klump     

When females should contest for food – testing hypotheses about resource density, distribution, size, and quality with Hanuman langurs (Presbytis entellus)

Predictions of ecological models on female social relationships (van Schaik 1989) and their links with food distribution and the potential competitive regime are used to analyze the feeding and spatial behavior, and resource density, size, distribution, and quality in a forest population of Hanuman langurs (Presbytis entellus). In contrast to other populations, and assumptions on folivorous primates, the females of this population exhibit a linear dominance hierarchy. The langurs concentrated their feeding on three key resources with a low density and clumped distribution. Two out of the three key resources contained significantly higher levels of extractable protein and soluble sugar than other food plants, indicating high spatial variability of food quality. Even the mature leaves of the most preferred food plant were about twice as nutritious as those from other food plants. Group spread was small and only a single high-quality resource was used at a time. Finally, even rich resources could accommodate only a subset of a group. These findings fit predictions made for the prevalence of within-group contest competition. Given the observed food distribution and phytochemical heterogeneity of mature foliage, even females of folivorous species should contest for food. The effect of female dominance rank on size and composition of feeding parties also agrees with this prediction. A comparison with data from another forest population, where female dominance relations are weakly developed, revealed a clear-cut difference in the use and abundance of resources. It is argued that between-population differences in female social relationships within a species may be viewed as adaptive responses to local habitat conditions. Received: 1 August 1997 / Accepted after revision: 7 December 1997     

Twilight fighting in the evening brown butterfly,<Emphasis Type="Italic"> Melanitis leda</Emphasis> (L.) (Nymphalidae): age and residency effects

Butterflies are conspicuous among animals that fight for mating opportunities because it is not clear how contest costs could accrue. Nevertheless, the bulk of research in this group suggests that contests are settled on the basis of asymmetries in fighting ability (the superior competitor hypothesis). There is also a consistent effect on contest escalation due to apparent 'confusion' over residency, which could result from resident butterflies playing different tactics than non-residents, or from a causal link between residency and fighting ability. I addressed these hypotheses by studying contest behavior in the crepuscular species Melanitis leda (L.) (Nymphalidae) over a 5-month period in tropical Australia. Males competed via conspicuous two-stage maneuvers of a form unique among butterflies. Prior residents won 77% of all contests. Non-resident males arrived and perched in occupied sites until challenged by incumbent residents, and the aggressiveness of these interlopers increased as a function of the time before they were detected. Contest winners also tended to be younger than losers, and contest escalation was negatively related to both the age of the losing male and the magnitude of the between-combatant age asymmetry. These results are consistent with the superior competitor hypothesis based upon age as a determinant (or correlate) of fighting ability, but also suggest residency has a unique influence on contest behavior. Further research is clearly required to unravel the effects of age, residency and other potential biophysical determinants of fighting ability in this species.     

Varying levels of female promiscuity in four <Emphasis Type="Italic">Apodemus</Emphasis> mice species

Sexual selection in most vertebrates is based on the evolution of fitness optimization strategies such as multiple-male mating (MMM). Several ecological correlates of MMM have been identified in bird and fish populations; however, only few studies have documented the effects of environmental change on promiscuity in mammals. In this study, the 127 pregnant females from four central European and ecologically diverse species of field mice (genus Apodemus) were studied to assess the role of ecological factors that may have shaped the evolution of particular mating systems. MMM was found in all analyzed species: in Apodemus uralensis and Apodemus flavicollis, up to two males could be identified as the fathers of a particular litter, while three males sired 9.1% of analyzed litters of Apodemus sylvaticus and 20.6% of Apodemus agrarius. Furthermore, there were obvious differences between species in relative testes size and the proportion of multiple sired litters during those seasons when the opportunity for multiple mating was high. The species with the smallest testes and the least promiscuous was A. uralensis (only 43.5% of multiple sired litters), while the species with the biggest testes and the most promiscuous was A. agrarius (69.2%). MMM was significantly associated with higher litter size in A. flavicollis, and the probability of MMM strongly increased with season in A. agrarius and with abundance in A. uralensis. These results indicate that ecological factors are associated with MMM rates in Apodemus field mice and more research is needed to fully understand the evolution of mating strategies at different levels of biological resolution.     

Competition,fluctuating asymmetry and sperm transfer in male gryllid crickets (<Emphasis Type="Italic">Gryllus bimaculatus</Emphasis> and <Emphasis Type="Italic">Gryllodes sigillatus</Emphasis>)

Sperm competition theory predicts that ejaculate expenditure by males should increase with the risk of competition from rivals. The tendency to modulate expenditure, however, may reflect various constraints acting on different individuals. We test this idea here by looking at the effects of competition and fluctuating asymmetry (FA) in two species of gryllid cricket. Male Gryllus bimaculatus and Gryllodes sigillatus modulated the sperm content of their spermatophores in relation to both apparent competition and FA in limb size. Males of both species increased the amount of sperm transferred when an apparent male competitor was present, sperm number increasing regardless of the species of apparent competitor in G. bimaculatus, but significantly more with a conspecific competitor in G. sigillatus. In addition, the relative size of the spermatophylax in G. sigillatus increased in the presence of apparent competition, while spermatophore transfer time decreased in G. bimaculatus. Sperm number showed opposite relationships with limb asymmetry in the two species, decreasing with increasing asymmetry in males in G. sigillatus, but increasing with asymmetry in both males and females in G. bimaculatus. G. bimaculatus, but not G. sigillatus, also transferred more sperm when paired with a larger female. Relationships between reproductive measures and limb asymmetry were significant only when competitors were present in G. bimaculatus, but were significant regardless of competition in G. sigillatus. The differences may reflect the additional burden of producing a spermatophylax in G. sigillatus.     

Host range and resistance to aspergillosis in three sea fan species from the Yucatan

Knowledge of host range and mechanisms of disease resistance is fundamental to predicting impacts and spread of marine diseases. Prevalence of signs of aspergillosis, caused by the terrestrial fungus Aspergillus sydowii, was assessed in the Yucatan among three species of sea fan: Gorgonia ventalina, G. cf. mariae, and G. flabellum. The Yucatan is unusual in that ranges of all three sea fan species overlap at many sites along a cline of increasing depth, allowing us to evaluate potential causes of differing prevalence among species. Signs of aspergillosis were observed in all the three species. However, the prevalence of infection in G. cf. mariae, a deep-water species, was consistently low even at sites where G. ventalina was common and had moderate levels of infection. Because G. cf. mariae is a relatively small-stature sea fan, we compared the prevalence of signs in G. cf. mariae to a subset of comparatively sized G. ventalina. G. ventalina had a significantly higher prevalence of aspergillosis, indicating that size does not explain the lower prevalence in G. cf. mariae. Prevalence of disease signs on the shallow-water G. flabellum was also significantly higher than G. cf. mariae, but did not differ from G. ventalina. To test the hypothesis that higher chemical resistance accounts for the low prevalence of disease in G. cf. mariae, we measured the response of A. sydowii in culture to antifungal extracts from each sea fan species. Significantly lower fungal growth rates on extracts of G. cf. mariae than G. ventalina support the hypothesis that G. cf. mariae is more chemically resistant to aspergillosis. When comparing sea fan disease across different regions of the Yucatan, we detected significantly higher prevalence in G. ventalina near Akumal than further north near Cozumel and Puerto Morelos. In Akumal, there was a strong positive correlation between sea fan size and disease, with the largest fans showing the highest prevalence and severity in all three species. In addition, prevalence of aspergillosis in G. ventalina was density-dependent in Ak.     

Causes and consequences of contest outcome: aggressiveness, dominance and growth in the sheepshead swordtail, Xiphophorus birchmanni

Although our understanding of how animal personality affects fitness is incomplete, one general hypothesis is that personality traits (e.g. boldness and aggressiveness) contribute to competitive ability. If so, then under resource limitation, personality differences will generate variation in life history traits crucial to fitness, like growth. Here, we test this idea using data from same-sex dyadic interaction trials of sheepshead swordtails (Xiphophorus birchmanni). In males, there was evidence of repeatable variation across a suite of agonistic contest behaviours, while repeatable opponent effects on focal behaviour were also detected. A single vector explains 80 % of the among-individual variance in multivariate phenotype and can be viewed as aggressiveness. We also find that aggressiveness predicts dominance—the repeatable tendency to win food in competition—and dominant individuals show faster post-trial weight gain (independently of initial size). In females, a dominance hierarchy predictive of weight gain was also found, but there was no evidence of variation in aggressiveness. While size often predicts contest outcome, our results show that individuals may sometimes grow larger because they are behaviourally dominant rather than vice versa. When resources are limited, personality traits such as aggression can influence growth, life history, and fitness through impacts on resource acquisition.     

Exaggerated male eye span influences contest outcome in stalk-eyed flies (Diopsidae)

Evolution of male weapons or status signals has been hypothesized to precede evolution of female mating preferences for those traits. We used staged male fights among three species of Malaysian stalk-eyed flies (Diptera: Diopsidae) to determine if elongated eye span, which is preferred by females in two sexually dimorphic species, influences contest outcome. Extreme sexual dimorphism, with large males possessing longer eye span than females, is shared by Cyrtodiopsis whitei and C. dalmanni. In contrast, C. quinqueguttata exhibits a more ancestral condition – short, sexually monomorphic eye stalks. Videotape analysis of 20-min paired contests revealed that males with larger eye span and body size won more fights in the dimorphic, but not monomorphic, species. To determine if males from the dimorphic species use eye span directly to resolve contests, we competed male C. dalmanni from lines that had undergone artificial selection for 30 generations to increase or decrease eye span. We found that eye span, independently of body size, determines contest outcome in selected-line males. Furthermore, in both dimorphic species, the average encounter duration declined as the eye span difference between contestants increased, as expected if males use eye span to assess opponent size. The number of encounters also increased with age in dimorphic, but not monomorphic, species. Selected-line males did not differ from outbred males in either fight duration or number of encounters. We conclude that exaggerated male eye stalks evolved to influence both competitive interactions and female mating preferences in these spectacular flies. Received: 20 July 1998 / Received in revised form: 2 February 1999 / Accepted: 12 March 1999     

The timing of calling,movement, and mating in the field crickets Gryllus veletis,G. pennsylvanicus,and G. integer

Summary The temporal relationships between mating, calling, and movement were studied in natural habitats at different densities in the field crickets, Gryllus veletis, G. pennsylvanicus and G. integer. There was variation between and within species, but males generally showed increased calling at dawn followed by a decline during the day. Matings were also more frequent at sunrise in most observations, and male and female movement declined near dawn. Female movement and sexual receptivity are closely related to levels of male calling and movement.     

The Capacity of Australia's Protected‐Area System to Represent Threatened Species

Abstract: The acquisition or designation of new protected areas is usually based on criteria for representation of different ecosystems or land‐cover classes, and it is unclear how well‐threatened species are conserved within protected‐area networks. Here, we assessed how Australia's terrestrial protected‐area system (89 million ha, 11.6% of the continent) overlaps with the geographic distributions of threatened species and compared this overlap against a model that randomly placed protected areas across the continent and a spatially efficient model that placed protected areas across the continent to maximize threatened species’ representation within the protected‐area estate. We defined the minimum area needed to conserve each species on the basis of the species’ range size. We found that although the current configuration of protected areas met targets for representation of a given percentage of species’ ranges better than a random selection of areas, 166 (12.6%) threatened species occurred entirely outside protected areas and target levels of protection were met for only 259 (19.6%) species. Critically endangered species were among those with the least protection; 12 (21.1%) species occurred entirely outside protected areas. Reptiles and plants were the most poorly represented taxonomic groups, and amphibians the best represented. Spatial prioritization analyses revealed that an efficient protected‐area system of the same size as the current protected‐area system (11.6% of the area of Australia) could meet representation targets for 1272 (93.3%) threatened species. Moreover, the results of these prioritization analyses showed that by protecting 17.8% of Australia, all threatened species could reach target levels of representation, assuming all current protected areas are retained. Although this amount of area theoretically could be protected, existing land uses and the finite resources available for conservation mean land acquisition may not be possible or even effective for the recovery of threatened species. The optimal use of resources must balance acquisition of new protected areas, where processes that threaten native species are mitigated by the change in ownership or on‐ground management jurisdiction, and management of threatened species inside and outside the existing protected‐area system.     

加拿大一枝黄花和小飞蓬叶浸提液对大豆幼苗生长的复合化感效应

加拿大一枝黄花(Solidago canadensis)和小飞蓬(Conyza canadensis)常共同入侵至同一农田生态系统。探究了两者对大豆幼苗生长的复合化感作用。加拿大一枝黄花叶浸提液显著抑制大豆幼苗地上生物量。而小飞蓬叶浸提液也明显抑制大豆幼苗地上生物量(未达到显著水平)。与对照处理相比,两者复合叶浸提液处理未显著影响大豆幼苗地上生物量。两者复合叶浸提液处理下大豆幼苗地上生物量显著大于加拿大一枝黄花叶浸提液单一处理,同时也大于小飞蓬叶浸提液单一处理(未达到显著水平)。两者叶浸提液单一处理均明显降低大豆幼苗株高(未达到显著水平)。与对照处理相比,两者复合叶浸提液处理未显著影响大豆幼苗株高。两者复合叶浸提液处理下大豆幼苗株高明显大于两者叶浸提液单一处理(未达到显著水平)。因此,两者叶浸提液对大豆幼苗的生长均具有一定程度的抑制效应,且加拿大一枝黄花叶浸提液对大豆幼苗产生的化感作用(尤其是地上生物量)明显大于小飞蓬。两者复合化感作用明显低于两者单一化感作用,尤其是两者对大豆幼苗地上生物量的复合化感作用显著低于加拿大一枝黄花的单一化感作用。所以,化感效应可能在两者共同入侵(即两者复合作用)进程中所起的贡献低于在两者单一入侵进程中所起的作用。     

Chemical mimicry of cuticular hydrocarbons – how does Eremostibes opacus gain access to breeding burrows of its host Parastizopus armaticeps (Coleoptera, Tenebrionidae)?

Summary. Breeding burrows of Parastizopus armaticeps armaticeps, a fossorial desert tenebrionid beetle, are cleptoparasitised by the closely related Eremostibes opacus. Gas chromatographic analyses show a high congruity of the cuticular hydrocarbons of both species. We compare these hydrocarbon patterns with those of four other Stizopina species and the Scaurini Herpiscius sommeri. In a bioassay, dummies treated with cuticular hydrocarbon extracts of E. opacus and the P. a. bifidus parasite E. bushmanicus were mostly ignored by P. a. armaticeps, whereas dummies with applied extracts of the remaining species were heavily attacked. We show that there is a correlation between agonistic behaviour of P. a. armaticeps towards the intruder and the chemical similarity of the cuticular hydrocarbons of the two species. Furthermore, we produced quantitatively modified hydrocarbon patterns of E. barbatus by changing the temperature at which this species was kept. The new 30 °C type was chemically similar to E. opacus, and was frequently ignored by P. a. armaticeps, whereas a reduction of the temperature to 20 °C only had minor effects on the hydrocarbon pattern. Furthermore, we show that the addition of one single component, heptacosane, to the cuticular hydrocarbon extract of E. opacus alters the host’s reaction. We discuss the role of cuticular hydrocarbons for the recognition of this host-parasite system and the relevance of quantitative characters in the hydrocarbon pattern for the discrimination of the host.     

Evolution of egg size and fertilisation efficiency in sea stars: large eggs are not fertilised more readily than small eggs in the genus<Emphasis Type="Italic"> Patiriella</Emphasis> (Echinodermata: Asteroidea)

Fertilisation kinetics theory suggests that, when sperm are limiting, the larger eggs of broadcast-spawning marine organisms ought to be fertilised more frequently than smaller eggs, because they provide a bigger target for searching sperm. Whilst this effect has been demonstrated within species, it is not known if this pattern holds among species. We tested whether a large difference in egg size between congeneric seastars with contrasting planktotrophic and lecithotrophic modes of development results in differences in the likelihood of eggs being fertilised in sperm-limiting situations. Measurement of egg sizes and sperm swimming speeds led to the prediction that the sperm–egg collision rate constant for Patiriella calcar (420-µm-diameter egg) should be nine times greater than for P. regularis (140-µm-diameter egg). Although the eggs of P. calcar should be fertilised at greater rates in low sperm concentrations, they were not. When gametes were allowed to mix for 10 s, the hypothesis that P. calcar eggs required less sperm than P. regularis to ensure 50% of eggs were fertilised was rejected. When gametes were mixed for 5 min, P. regularis eggs were more frequently fertilised, but the difference was not statistically significant. We conclude there must be a difference between these species in the likelihood that when a sperm finds a conspecific egg it can successfully fertilise. This apparent uncoupling of egg size and likelihood of fertilisation suggests that fertilisation is not a major constraint on the evolution of egg size in these seastars.Communicated by G.F. Humphrey, Sydney     

Biological traits of three closely related species of <Emphasis Type="Italic">Pycnoclavella</Emphasis> (Ascidiacea) in the Western Mediterranean

The reproductive cycles and abundance of the sympatric colonial ascidians Pycnoclavella brava, Pycnoclavella aurilucens and Pycnoclavella communis from two Northwestern Mediterranean sites over a period of 2.5 years are reported. The species showed some differences in their biological patterns. Gonad maturation and larval brooding took place during autumn and early winter for P. communis, during spring for P. aurilucens and during late winter and spring for P. brava. Summer was found to be the unfavorable season for all species, and an aestivation period occurred in P. communis and P. brava. Maximum abundance for the three species was observed during winter. Growth rates of the most abundant species, P. communis, were also monitored and found to peak during the initial phases of reactivation after aestivation. Growth rates decreased afterwards, becoming negative as water temperature started to increase after the winter minima. Reproductive activity and growth rates for P. communis displayed a temporal lag that suggested partitioning of resources to either reproduction or growth in this species.     

Size and spatial arrangement of home range of checkered snapper <Emphasis Type="Italic">Lutjanus decussatus</Emphasis> (Lutjanidae) in an Okinawan coral reef determined using a portable GPS receiver

Although snappers (Lutjanidae) are commercially important fishery resources in tropical and subtropical waters, their home range size and its spatial arrangement have not been sufficiently clarified. In the present study, the size and spatial arrangement of the home range of the reef-associated checkered snapper Lutjanus decussatus (Lutjanidae) were investigated by use of a portable GPS receiver. In a 120 m × 120 m quadrat established on a section of the fringing reef of Ishigaki Island, 29 individuals of the species were identified by their color pattern and individuals were divided into five arbitrary size classes (class 1 <10 cm TL ≤ class 2 <15 cm TL ≤ class 3 <20 cm TL ≤ class 4 <25 cm TL ≤ class 5). Fish tracking by snorkeling was conducted with the portable GPS receiver. Site fidelity of this species was high. Home range size ranged from 93.0 to 3638.4 m², and there was a significant positive correlation between the home range size and fish total length. Home ranges of the same-sized individuals abutted each other (8.8% area overlap), whereas those of different-sized individuals overlapped (44.0% area overlap). Agonistic behavior (attack and agonistic display) was more frequently found among same-sized individuals (times of agonistic behavior/times of all encounters × 100 = 71.3%), whereas such agonistic behavior was rarely found among different-sized individuals (times of agonistic behavior/times of all encounters × 100 = 6.9%). These results suggest that home ranges of Lutjanus decussatus can be regarded as territories against same-sized individuals, but not different-sized individuals. The usefulness of the fish tracking by snorkeling using a portable GPS receiver for home range size estimation and the function of the overlapping territory of the species are discussed.     

New repellent semiochemicals for three species of <Emphasis Type="Italic">Dendroctonus</Emphasis> (Coleoptera: Scolytidae)

Summary. Nine compounds identified from captured volatiles of the Douglas-fir beetle, Dendroctonus pseudotsugae, the mountain pine beetle, D. ponderosae, and the spruce beetle, D. rufipennis, that elicited antennal responses in males and females of one or more of these species were tested in the field to determine behavioural activity. 1-Octen-3-ol, found in the volatiles of females of all three species decreased the response of male and female coastal and male interior D. pseudotsugae and both sexes of D. ponderosae to their aggregation pheromones. Acetophenone, identified in the volatiles of females of all three species, significantly decreased the response of interior female D. pseudotsugae. trans-Verbenol, a potent aggregation pheromone of D. ponderosae, decreased the response of both sexes of D. pseudotsugae, while 3-methyl-2-cyclohexen-1- one (MCH), the antiaggregation pheromone of D. pseudotsugae and D. rufipennis decreased the response of both sexes of D. ponderosae. While it has been demonstrated that semiochemical mediated interspecific communication occurs among bark beetles infesting the same host, this study demonstrates that beetles can perceive signals emitted by heterospecifics attacking nonhosts and can potentially use them to avoid attacking the wrong species of conifer.     

Intraspecific competition in<Emphasis Type="Italic"> Fucus serratus</Emphasis> and<Emphasis Type="Italic"> F</Emphasis>.<Emphasis Type="Italic"> evanescens</Emphasis> (Phaeophyceae: Fucales) germlings: effects of settlement density,nutrient concentration,and temperature

Survival and growth of early post-settlement stages are critical for the development of seaweed populations. Fucoid germlings commonly settle in dense monospecific aggregates, where intraspecific competition and environmental variables (e.g. nutrient concentration and temperature) may affect survival and growth. Using factorial experiments, we determined the effects of settlement density (~10, ~50 and ~250 germlings cm^–2), nutrient enrichment (from ~10 to ~40 µM N and from ~0.5 to ~2.5 µM P), and temperature (7°C and 17°C) on Fucus serratus and F. evanescens germlings in laboratory cultures over 3 months. Settlement density, nutrient concentration and temperature interactively affected growth of germlings, and the magnitude of this interaction varied between the two species. This represents the first record of such factorial interactions in Fucus spp. germlings. Intraspecific competition, estimated as the relative reduction in germling growth and survival from low to high densities, increased with decreasing nutrient concentration and increasing temperature in both species. While temperature and nutrient concentration had little effect on germling size distributions, size inequality and skewness generally increased with germling density, indicating that a few large individuals gained dominance and suppressed many smaller ones at high density. Self-thinning increased with settlement density and depended on nutrient concentration and species at high density. At high density, self-thinning increased with decreasing nutrient levels in F. evanescens, but not in F. serratus. At low density, nutrient enrichment increased germling growth in F. evanescens, but not in F. serratus, whereas growth in both species was stimulated by nutrient enrichment at higher densities. These results suggest that germling growth and self-thinning are more sensitive to variation in nutrient concentration in F. evanescens than in F. serratus. The potential implications of our findings for the understanding of eutrophication-related abundance changes in both species in southern Norway are discussed.Communicated by L. Hagerman, Helsingør     

Fighting behavior in hermit crabs: the combined effect of resource-holding potential and resource value in Pagurus longicarpus

A study was conducted to determine whether asymmetries in both resource-holding potential (RHP) and resource value (RV) influence dominance and fighting behavior in the hermit crab Pagurus longicarpus. A total of 120 groups of three crabs were observed for 10 min in four experiments that tested all diverse combinations of equal/different RHP (i.e. ‘body size’) and equal/different RV (i.e. ‘shell size’ and ‘shell quality’). In a fifth experiment, dominant and subordinate individuals of the same size category (26 groups) were forced to enter shells of opposite quality than those previously occupied, and then the behavior of the reconstituted original groups was observed for additional 10 min. As expected, crabs in lower quality shells were more willing to initiate and to escalate fights. However, their attacks were directed to any crab of the group, independently of the defender's shell quality, and the fight duration did not vary with the different value of the resources at stake. This may indicate that P. longicarpus is unable to assess the quality of the shells available in its social environment but bases its tactical decisions during fights solely on its own resource. This suggestion was confirmed by the change in the fighting behavior of crabs whose shell quality was experimentally altered. This manipulation induced an overall increase in the intensity of aggression, drastic modification of crab behavior, and inversion of the hierarchy even though these crabs have had previous experiences of wins/losses and were familiar to the other members of the group. In this species, large crab size and/or the occupancy of adequate (and oversized) shells appeared to be the most likely determinant of contest resolution. Individuals seemed to retain a memory of the previously held resource and behaved accordingly.     

Size-related aspects of arm damage, tissue mechanics, and autotomy in the starfish Asterias rubens

 Arm damage is a widely reported but superficially investigated aspect of the biology of the starfish Asterias rubens L. In the present study, the incidence of arm damage was surveyed in populations of A.  rubens at two sites in the Firth of Clyde, Scotland, and three sites in Gullmarsfjorden, Sweden. The mean (±SD) incidence across all sites of individuals with basal arm damage (resulting from detachment at the basal autotomy plane) was 19.69 ± 8.86%, the incidence of those with distal arm damage (resulting from amputation at more distal levels) was 7.74 ± 10.01%. The mean incidence of arms with basal damage was 5.28 ± 4.12%, of those with distal damage 1.83 ± 2.45%. There was a significant negative correlation between size and the incidence of basal damage at all but one site, but no significant correlation between size and distal damage at any site. Mechanical tests on specimens of the aboral body wall from the basal region of the arm (which included the autotomy plane) and from a more distal region revealed that with increasing body size there was a significant increase in yield stress, ultimate stress and Young's modulus (stiffness) but no significant change in yield strain and ultimate strain. There was no significant difference between the relationships for basal and distal specimens. It is hypothesised that in larger individuals increased mechanical toughness replaces autotomy as an effective antipredator strategy. Using two methods to induce autotomy, a significant positive correlation between size and the delay between the onset of stimulation and arm detachment was found; this may represent a size-related decline in the efficiency of the autotomy mechanism through the relaxation of selection pressure. Since size is an unreliable indicator of age in A.  rubens, the trends identified herein can be interpreted only tentatively as age-associated phenomena. Received: 4 September 1999 / Accepted: 17 February 2000