Multimodal signals,imperfect information,and identification of sex in red-backed salamanders (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>)

Theoretical treatments and empirical studies both suggest that signals that occur in multiple sensory modes have superior detectabilities, discriminabilities, and memorabilities. There is also an intuitive link between signal detectability, discriminability, and memorability and the quality of information that is transferred via the signaler. We investigated the role of information quality and sensory modality in the sexual identification of intruding conspecifics by territorial male red-backed salamanders (Plethodon cinereus). Our results imply that these salamanders are able to discriminate between intruding male and female conspecifics (based on the allocation of aggression) in bimodal scenarios (vision and olfaction) even under situations in which the information available about the intruders sexual identities is reduced in quality. In unimodal scenarios (olfaction only), male residents exhibited heightened levels of aggression toward male secretions and reduced levels of aggression toward female secretions. In unimodal scenarios where information pertaining to the sex of the stimulus was reduced in quality, male residents displayed intermediate levels of aggression relative to the responses that male and female stimuli received respectively. Although resident males touched bimodal stimuli significantly more than they touched unimodal stimuli, we were unable to find support for the notion that sensory modality greatly influences how male resident red-backed salamanders allocate aggression toward intruding male versus female stimuli.Communicated by H. Kokko     

Rapid onset of mate quality assessment via chemical signals in a woodland salamander (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>)

In the competition to acquire mates, environmental factors can be important in determining the relative quality of an individual. These aspects of quality are often conveyed through signals used for mate assessment by the most energetically invested sex. In red-backed salamanders, Plethodon cinereus, chemical signals communicate a surprising amount of information, much of which is dependent on the current condition of the sender. One such condition-dependent aspect of information conveyed via chemicals is the quality of the sender’s diet, but it is unclear as to whether this information is actively advertised by the sender (i.e., a signal) or simply inferred from food-derived odors (i.e., a cue). The amount of time on different diets required for changes in signaling is also unknown. I examined how quickly gravid female salamanders could detect a difference between the scents of males on high- vs. low-quality diets without fecal cues. The amount of protein present in two known signaling glands (the mental and postcloacal glands) was also measured after experimental feeding. Gravid females were able to infer the differences in male diet quality after only 1 week. Females also responded to the male scents more quickly after differential feeding had begun. High-quality males had significantly more protein present in both signaling glands than low-quality males. This scenario highlights the ongoing interplay between the quality of an individual and its environment, with males actually advertising the status of this relationship as an honest signal for mate assessment.     

Genetic analysis of tuna populations,<Emphasis Type="Italic"> Thunnus thynnus thynnus</Emphasis> and<Emphasis Type="Italic"> T</Emphasis>.<Emphasis Type="Italic"> alalunga</Emphasis>

The genetic population structures of Atlantic northern bluefin tuna ( Thunnus thynnus thynnus) and albacore ( T. alalunga) were examined using allozyme analysis. A total of 822 Atlantic northern bluefin tuna from 18 different samples (16 Mediterranean, 1 East Atlantic, 1 West Atlantic) and 188 albacore from 5 samples (4 Mediterranean, 1 East Atlantic) were surveyed for genetic variation in 37 loci. Polymorphism and heterozygosity reveal a moderate level of genetic variability, with only two highly polymorphic loci in both Atlantic northern bluefin tuna ( FH* and SOD- 1*) and albacore ( GPI- 3* and XDH*). The level of population differentiation found for Atlantic northern bluefin tuna and albacore fits the pattern that has generally been observed in tunas, with genetic differences on a broad rather than a more local scale. For Atlantic northern bluefin tuna, no spatial or temporal genetic heterogeneity was observed within the Mediterranean Sea or between the East Atlantic and Mediterranean, indicating the existence of a single genetic grouping on the eastern side of the Atlantic Ocean. Very limited genetic differentiation was found between West Atlantic and East Atlantic/Mediterranean northern bluefin tuna, mainly due to an inversion of SOD- 1* allele frequencies. Regarding albacore, no genetic heterogeneity was observed within the Mediterranean Sea or between Mediterranean and Azores samples, suggesting the existence of a single gene pool in this area.     

Socially facilitated antipredator behavior by ringed salamanders (<Emphasis Type="Italic">Ambystoma annulatum</Emphasis>)

Many aspects of animal behavior can be socially facilitated, including foraging behavior, exploration behavior, and antipredator behavior. Although larvae of the ringed salamander (Ambystoma annulatum) are not gregarious, they can live in high densities and face intense predation pressure during a short period following hatching. In a predator-recognition experiment, we found that these salamanders responded to chemical cues from dragonfly nymphs (Family: Libellulidae) with appropriate antipredator behavior (decreased activity), and this response was absent when salamanders were exposed to chemical cues from nonpredatory mayfly nymphs (Family: Heptageniidae). In a second experiment, we tested whether antipredator behavior in response to chemical cues of dragonflies could be socially facilitated by larval ringed salamanders. We placed an “observer” salamander into a central arena with four “demonstrator” salamanders behind clear barriers around an arena. The barriers ensured that chemical cues would not be detected by the observer. When demonstrators were exposed to chemical cues from dragonflies, the data were consistent with the hypothesis that both demonstrators and observers decreased activity relative to a blank control. Our results provide evidence that social facilitation can occur in larval ringed salamanders, a nonsocial species.     

Cofoundress relatedness and group productivity in colonies of social <Emphasis Type="Italic">Dunatothrips</Emphasis> (Insecta: Thysanoptera) on Australian <Emphasis Type="Italic">Acacia</Emphasis>

Facultative joint colony founding by social insects provides opportunities to analyze the roles of genetic and ecological factors in the evolution of cooperation. Although cooperative nesting is observed in range of social insect taxa, the most detailed studies of this behavior have been conducted with Hymenoptera (ants, bees, and wasps). Here, we show that foundress associations in the haplodiploid social thrips Dunatothrips aneurae (Insecta: Thysanoptera) are most often comprised of close relatives (sisters), though groups with unrelated foundresses are also found. Associations among relatives appear to be facilitated by limited female dispersal, which results in viscous population structure. In addition, we found that per capita productivity declined with increasing group size, sex ratios were female-biased, and some female offspring apparently remained in their natal domicile for some time following eclosion. D. aneurae thus exhibits a suite of similarities with eusocial Hymenoptera, providing evidence for the convergent evolution of associated social and life-history traits in Hymenoptera and Thysanoptera.     

Auditory defence in the peacock butterfly (<Emphasis Type="Italic">Inachis io</Emphasis>) against mice (<Emphasis Type="Italic">Apodemus flavicollis</Emphasis> and <Emphasis Type="Italic">A. sylvaticus</Emphasis>)

Morphological and behavioural traits can serve as anti-predator defence either by reducing detection or recognition risks, or by thwarting initiated attacks. The latter defence is secondary and often involves a ‘startle display’ comprising a sudden release of signals targeting more than one sensory modality. A suggested candidate for employing a multimodal defence is the peacock butterfly, Inachis io, which, by wing-flicking suddenly, produces sonic and ultrasonic sounds and displays four large eyespots when attacked. The eyespots make small birds retreat, but whether the sounds produced thwart predator attacks is largely unknown. Peacocks hibernate as adults in dark wintering sites and employ their secondary defence upon encounter with small rodent predators during this period. In this study, we staged predator–prey encounters in complete darkness in the laboratory between wild mice, Apodemus flavicollis and Apodemus sylvaticus, and peacocks which had their sound production intact or disabled. Results show that mice were more likely to flee from sound-producing butterflies than from butterflies which had their sound production disabled. Our study presents experimental evidence that the peacock butterfly truly employs a multimodal defence with different traits targeting different predator groups; the eyespots target birds and the sound production targets small rodent predators.     

Histological investigation of the reproductive cycles of the limpets <Emphasis Type="Italic">Patella </Emphasis><Emphasis Type="Italic">vulgata</Emphasis> and <Emphasis Type="Italic">Patella ulyssiponensis</Emphasis>

This study devised a staging system for, and monitored, the gonad development of the limpet species Patella vulgata and Patella ulyssiponensis on the South West coast of Ireland using histological techniques. Maturation began in the males of both species in January and in the females it began in March. There was no statistical difference in gonad development between sexes and between species. Spawning in the male P. vulgata occurred from September to December 2003 and in September and October 2004. In female P. vulgata spawning occurred from October to December 2003, no spawning of females was observed in 2004. In male P. ulyssiponensis spawning occurred in November and December 2003 and from September 2004 to December 2004. Spawning was observed from November 2003 to January 2004 and in September 2004 in female P. ulyssiponensis. Sex ratios also varied between the species and between months sampled. Nevertheless more males were observed in both species.     

Comparative life-history study on sympatric hyperiid amphipods (<Emphasis Type="Italic">Themisto pacifica</Emphasis> and <Emphasis Type="Italic">T</Emphasis>. <Emphasis Type="Italic">japonica</Emphasis>) in the Oyashio region,western North Pacific

Life-history features of the sympatric amphipods Themisto pacifica and T. japonica in the western North Pacific were analyzed based on seasonal field samples collected from July 1996 through July 1998, and data from laboratory rearing experiments. T. pacfica occurred throughout the year, with populations peaking from spring to summer. In contrast, T. japonica were rare from autumn to early winter, but became abundant in late winter to spring. Mature T. pacifica females and juveniles occurred together throughout the year, indicating year-round reproduction. Mature T. japonica females were observed only in spring, and juveniles occurred irregularly in small numbers, suggesting limited, early-spring reproduction in this study area. Size composition analysis of T. pacifica identified a total of eight cohorts over the 2 years of the study. Due to the smaller sample size and rarity of mature females (>9.6 mm) and males (>7.1 mm), cohort analyses of T. japonica were not comparable. Laboratory rearing of specimens at 2°C, 5°C, 8°C and 12°C revealed that a linear equation best expressed body length growth by T. pacifica, while a logistic equation best expressed body length growth by T. japoncia. Combining these laboratory-derived growth patterns with maturity sizes of wild specimens, the minimum and maximum generation times of females at a temperature range of 2–12°C were computed as 32 days (12°C) and 224 days (2°C), respectively, for T. pacifica, and 66 days (12°C) and 358 days (2°C), respectively, for T. japonica. The numbers of eggs or juveniles in females marsupia increased with female body length and ranged from 23 to 64 for T. pacifica and from 152 to 601 for T. japonica. Taking into account the number of mature female instars, lifetime fecundities were estimated as 342 eggs for T. pacifica and 1195 eggs for T. japonica. Possible mechanisms for the coexistence of these two amphipods in the Oyashio region are also discussed.Communicated by O. Kinne, Oldendorf/Luhe     

Classification of juvenile rockfish,<Emphasis Type="Italic"> Sebastes inermis</Emphasis>, to<Emphasis Type="Italic"> Zostera</Emphasis> and<Emphasis Type="Italic"> Sargassum</Emphasis> beds,using the macrostructure and chemistry of otoliths

Young-of-the-year (YOY) Sebastes inermis use Zostera and Sargassum beds as nursery grounds, although it is not known which habitat YOY prefer. In this study, YOY S. inermis were accurately assigned to Zostera or Sargassum beds by two approaches: the width and length of the otolith nucleus and the composition of trace elements in otoliths. The otolith nucleus was initially opaque and then showed a marked shift to hyaline deposition once YOY settled in the nursery grounds. The first hyaline zone (FHZ) was deposited earlier in Zostera beds (from mid-May to early June) than in Sargassum beds (around mid-summer). Likewise, irrespective of settlement year, the FHZ was formed at both significantly younger ages and shorter back-calculated sizes (total length, TL) in the Zostera bed (overall mean: 131±3 days; 2.5±1.7 mm TL) than in the Sargassum bed (overall mean: 158±12 days; 61.3±1.00 mm TL). YOY collected in the Zostera bed were born earlier (mainly in January) than YOY from the Sargassum bed (mainly in February). In addition, a significant linear relationship was found between the age at formation of the FHZ and nucleus dimensions, suggesting that nucleus dimensions were a reliable macroscopic indicator of the time of formation of the FHZ and, consequently, also an indicator of the nursery where YOY grew. Linear discriminant function analysis (LDFA) based on the width and length of the otolith nucleus could distinguish juveniles from Zostera (88–96%) and Sargassum (96–97%) beds with a high degree of accuracy. In the other approach, six detectable trace elements (Li, Mn, Ni, Cu, Zn, and Ba) in otoliths of YOY collected in the nursery grounds were measured by high-resolution, inductively coupled mass spectrometry. LDFA based on the trace elemental composition separated YOY from three nurseries with 100% of accuracy. Findings suggest that both the trace elemental composition and nucleus dimensions of otoliths can be used as natural tags of the nursery grounds of S. inermis, offering a considerable potential for answering questions on habitat use and the contribution of nursery grounds to the adult stock.Communicated by T. Ikeda, Hakodate     

Morphology and growth of the deep-water gorgonians <Emphasis Type="Italic">Primnoa resedaeformis</Emphasis> and <Emphasis Type="Italic">Paragorgia arborea</Emphasis>

The morphology of the gorgonian corals Paragorgia arborea and Primnoa resedaeformis was studied from video records and colonies collected from different locations in Atlantic Canada, at depths between 200 and 600 m. Growth was studied by relating colony height to age (number of growth rings) in P. resedaeformis, and from a photographic time-series of a P. arborea colony in a Norwegian fjord. The highest P. resedaeformis and P. arborea colonies were 86 and 180 cm, respectively. The height of P. arborea seemed to be restricted by the size of the boulder it was attached to. When the coral exceeds a critical height (approximately twice the stone size), the drag of strong currents can turn the coral and its substrate over. No limiting factors for the height of P. resedaeformis colonies were identified. P. arborea occurred in three colour varieties: red, salmon red, and white. The red and white contributed 41% to the population each, while 18% of the colonies were salmon red. On average the salmon red P. arborea were taller than the red and white. P. arborea colonies >50 cm were mainly concave fan shaped. The orientation of these indicated a near-bottom current pattern similar to what is known from previous current measurements in the area. P. resedaeformis occurred mainly on the up-current side of boulders, but its bushy morphology does not indicate influence by unidirectional current to the same degree as P. arborea. The different height, morphology, and position on boulders of the two species indicate that they utilize different food sources. P. resedaeformis seems to be adapted to a near-bottom environment with turbulent currents, whereas P. arborea utilize uni- or bidirectional currents higher above bottom by developing planar colonies perpendicular to the current. The oldest P. resedaeformis colony was 61 years. The relationship between height and age indicated an average growth of 1.7 cm year^–1 for P. resedaeformis. X-ray images of skeletal sections of P. arborea showed clear growth bands with a maximum band width of 1.3 cm. It is not clear what time scales these bands represent, and they could therefore not be used for indicating age. The limited previously reported data on age and growth of P. arborea indicate an average growth rate of 1 cm year^–1. This gives an age of about 180 years for the largest colony in this study. The time-series photographs, however, indicated a much higher growth rate (varying between 2 and 6 cm year^–1 within the colony), which may be more representative for colonies of an intermediate size.Communicated by R.J. Thompson, St. Johns     

Geographical genetic structure and phylogeography of the <Emphasis Type="Italic">Sargassum horneri</Emphasis>/<Emphasis Type="Italic">filicinum</Emphasis> complex in Japan,based on the mitochondrial <Emphasis Type="Italic">cox3</Emphasis> haplotype

The genetic structure and phylogeography of the brown seaweed Sargassum horneri/filicinum complex in Japan were studied based on the mitochondrial cox3 haplotype. The cox3 haplotypes found were divided into three clades in a statistical parsimony network, among which there were large numbers of steps. Contrary to the reported large amount of drifting S. horneri along the Japanese coast, the three clades were dividedly distributed on the Japanese coast: the northern Pacific, the central Pacific, and western Japan. The western Japan S. horneri had haplotypes that were phylogenetically closer to those of S. filicinum than to the northern and central Pacific S. horneri populations. The S. filicinum populations were included within the western Japan clade and grouped together with the S. horneri samples from western Japan. Taken together with the unstable morphological diagnosis, this result suggests that S. filicinum should be reduced into a synonymy of S. horneri. The TMRCA analysis suggested that the divergence time of each clade may go back to the last interglacial period and a skyline plot suggested that the last glacial maximum had only a small effect on the population size of S. horneri. The geographic subdivision of the three groups, in spite of a large amount of drifting mats, suggests a limited contribution of drifting mats to gene flow on a large geographic scale. On a small geographic scale, a small number of haplotypes were shared between S. horneri-type and S. filicinum-type populations. This result suggests that populations of these two types are partially, though not completely, isolated from each other, possibly by selfing in S. filicinum-type populations or by a difference in peak reproduction.     

Predation on egg capsules of <Emphasis Type="Italic">Zidona</Emphasis><Emphasis Type="Italic">dufresnei</Emphasis> (Volutidae): ecological implications

Among the diverse patterns of energy allocation to the offspring of gastropods, the presence of egg capsules to protect embryos is common. Females of the edible snail Zidona dufresnei attach egg capsules to hard substrates in shallow Argentine Patagonian waters (40°45′S, 64°56′W) during spring-summer. Embryonic development takes about 30 days at 22°C. In this study, three likely capsule predator species and the marks left by each on egg capsule walls were identified in laboratory experiments in February 2010. Abundances of predators and egg capsules with evidence of predation were assessed in the field in the summers of 2010 and 2011. Under laboratory conditions (N = 10 replicates per treatment and control), the predation rate by the chiton Chaetopleura isabellei was the highest (up to 90%), followed by the gastropod Tegula patagonica and the crab Neohelice granulata (~20% each). Nearly 60% of 41 capsules found in the field showed signs of predation. According to the marks identified in the laboratory, C. isabellei was responsible for 79% of this predation, and T. patagonica for the rest. Predation appears to be important during the encapsulated early life and could be an agent for selecting for resistant capsule walls and a relatively shorter development time.     

Insights on common dolphin (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) social organization from genetic analysis of a mass-stranded pod

Compared to terrestrial mammals, little is known of cetacean social systems as they are generally less accessible to behavioral investigations due to their aquatic environment. The present study investigates group structure of the pelagic common dolphin, Delphinus delphis, using genetic markers. Tissue samples from 52 individuals representing a recent live mass-stranding event were compared to 42 single strandings taken from presumably different groups. The mass-stranding event occurred in 2002 on the French coast of the English Channel, whereas the single strandings were collected between 1993 and 2003 along the western coast of France (Bay of Biscay and English Channel). Analysis of mitochondrial DNA control region sequences indicated that genetic variability within the mass-stranded pod was similar to variability observed in single strandings. The mass-stranded group was composed of 41 different mitochondrial haplotypes or matrilines while the single strandings revealed 29 different haplotypes. Analysis of 11 microsatellite loci revealed that average relatedness of the mass-stranded pod was not different from average relatedness among all single strandings suggesting that individuals within the group had no closer kin relationships than animals taken from presumably different groups. These results do not support a matriarchal system and suggest that common dolphins constituting a pod are not necessarily genetically related.     

Patterns of gamete incompatibility between the blue mussels<Emphasis Type="Italic"> Mytilus edulis</Emphasis> and<Emphasis Type="Italic"> M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis>

Previous research on gametic incompatibility in marine invertebrates suggests that for highly dispersive marine invertebrate species, barriers to fertilization among closely related taxa are often incomplete and sometimes asymmetric. The nature of these barriers can dramatically affect the patterns of gene flow and genetic differentiation between species, and thus speciation. Blue mussels, in the genus Mytilus, are genetically distinct in allopatry yet hybrids are present wherever any two species within the group co-occur. The present study sampled M. edulis (L.) and M. trossulus (Gould) in May and June 2001 from the East Bay section of Cobscook Bay, Maine, USA (latitude 44°56′30″N; longitude 67°07′50″W), where the two species are sympatric. Gamete incompatibility was investigated in a series of laboratory fertilizations carried out in July 2001. The proportion of fertilized eggs typically exceeded 80% at sperm concentrations of 10³–10⁴ ml^?1 among intraspecific matings (n=18), but was <30% even at sperm concentrations in excess of 10⁵–10⁶ ml^?1 for interspecific matings (n=13). Further analysis indicated that approximately 100- to 700-fold higher sperm concentrations were required to achieve 20% fertilization in interspecific matings relative to intraspecific matings, indicating strong barriers to interspecific fertilization. The proportion of fertilized eggs did not follow this general pattern in all matings, however. The eggs from two (out of five) M. edulis females were almost as receptive to M. trossulus sperm as they were to M. edulis sperm. In contrast, the eggs from all M. trossulus females (n=3) were unreceptive to M. edulis sperm, suggesting that fertilization barriers between these species may be asymmetric. Given the experimental design employed in this study, the results are also consistent with a strong maternal or egg effect on the level of interspecific gamete compatibility in M. edulis.     

Feeding selectivities of the marine cladocerans<Emphasis Type="Italic"> Penilia avirostris</Emphasis>,<Emphasis Type="Italic"> Podon intermedius</Emphasis> and<Emphasis Type="Italic"> Evadne nordmanni</Emphasis>

We conducted grazing experiments with the three marine cladoceran genera Penilia, Podon and Evadne, with Penilia avirostris feeding on plankton communities from Blanes Bay (NW Mediterranean, Spain), covering a wide range of food concentrations (0.02–8.8 mm³ l^–1, plankton assemblages grown in mesocosms at different nutrient levels), and with Podon intermedius and Evadne nordmanni feeding on the plankton community found in summer in Hopavågen Fjord (NE Atlantic, Norway, 0.4 mm³ l^–1). P. avirostris and P. intermedius showed bell-shaped grazing spectra. Both species reached highest grazing coefficients at similar food sizes, i.e. when the food organisms ranged between 15 and 70 µm and between 7.5 and 70 µm at their longest linear extensions, respectively. E. nordmanni preferred organisms of around 125 µm, but also showed high grazing coefficients for particles of around 10 µm, while grazing coefficients for intermediate food sizes were low. Lower size limits were >2.5 µm, for all cladocerans. P. avirostris showed upper food size limits of 100 µm length (longest linear extension) and of 37.5 µm particle width. Upper size limits for P. intermedius were 135 µm long and 60 µm wide; those for E. nordmanni were 210 µm long and 60 µm wide. Effective food concentration (EFC) followed a domed curve with increasing nutrient enrichment for P. avirostris; maximum values were at intermediate enrichment levels. The EFC was significantly higher for P. intermedius than for E. nordmanni. With increasing food concentrations, the clearance rates of P. avirostris showed a curvilinear response, with a narrow modal range; ingestion rates indicated a rectilinear functional response. Mean clearance rates of P. avirostris, P. intermedius and E. nordmanni were 25.5, 18.0 and 19.3 ml ind.^–1 day^–1, respectively. Ingestion rates at similar food concentrations (0.4 mm³ l^–1) were 0.6, 0.8 and 0.9 g C ind.^–1 day^–1.Communicated by O. Kinne, Oldendorf/Luhe     

Heart rate reflects osmostic stress levels in two introduced colonial ascidians <Emphasis Type="Italic">Botryllus schlosseri</Emphasis> and <Emphasis Type="Italic">Botrylloides</Emphasis><Emphasis Type="Italic">violaceus</Emphasis>

The influence of abiotic factors on the establishment and success of invasive species is often difficult to determine for most marine ecosystems. However, examining this relationship is critical for predicting the spread of invasive species and predicting which habitats will be most vulnerable to invasion. Here we examine the mortality and physiological sensitivity to salinity of adult colonies of the colonial ascidians Botryllus schlosseri and Botrylloides violaceus. Adult colonies of each species were exposed to abrupt changes in salinity (5, 10, 15, 20, 25, 30 psu) in the laboratory. Salinity ranges used in the laboratory corresponded with those of the field distributions of B. violaceus and B. schlosseri in the Great Bay Estuary, NH. Heart rate was used as a proxy for health to assess the condition of individual colonies. Heart rates were monitored daily for approximately 2 weeks. Results revealed that both species experienced 100% mortality after 1 day at 5 psu and that their heart rates declined with decreasing salinity. Heart rates of B. schlosseri remained consistent between 15 and 30 psu and slowed at 10 psu. Heart rates of B. violaceus remained constant between 20 and 30 psu, but slowed at 15 psu. These laboratory results corresponded to the distribution of these species in estuaries, indicating salinity is a key factor in the distribution and dominance of B. schlosseri and B. violaceus among coastal and estuarine sites. Furthermore, physiological differences to salinity were found between colonies of B. schlosseri in the Venetian Lagoon and colonies in Portsmouth Harbor, suggesting adaptation to environmental variables.     

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Cooperation in animal social groups may be limited by the threat of free riding, the potential for individuals to reap the benefits of other individuals actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other groups territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.Communicated by P. Kappeler     

Disinfection efficacy on cyst viability of<Emphasis Type="Italic"> Artemia franciscana</Emphasis> (Crustacea),<Emphasis Type="Italic"> Hexarthra fennica</Emphasis> (Rotifera) and<Emphasis Type="Italic"> Fabrea salina</Emphasis> (Ciliophora)

Resting stages (cysts) of Artemia franciscana, Hexarthra fennica and Fabrea salina were exposed for two periods (acute and chronic exposures) to different concentrations of five aquaculture disinfectants (formalin, sodium hypochlorite, potassium permanganate, organic iodine, copper sulphate). The effects of chemical treatments were ranked according to their action on cyst hatching: total inhibition (no cyst hatching), heavy inhibition (significant decrease of cyst hatching), alteration of hatching synchrony, and occurrence of anomalous hatching (death of emerging individuals or birth of malformed organisms). Resting-stage viability was not completely impaired by disinfectant exposures at the ordinary doses used in aquaculture, so that usual treatments are possibly ineffective against the protist and metazoan cysts potentially present in breeding systems. On the other hand, the high resistance shown by cysts suggests the possibility of using these chemicals in live-feed disinfection.