Isolation and characterization of microsatellite markers for the White Cloud Mountain minnow (Tanichthys albonubes) in wild and cultured populations

We developed 12 microsatellite loci for the endangered minnow species, Tanichthys albonubes, using PCR-based isolation of microsatellite arrays. These new markers were tested in 26 individuals from a wild population collected from Guangzhou in China and 26 individuals from a cultured strain. The number of alleles ranged from two to nine and the expected heterozygosity from 0.177 to 0.853. The wild population had significantly higher allelic richness than the cultured strain, with a mean allelic richness of 5.52 (range = 3.69-8.64) and 3.13 (range = 1.99-5.73) for the wild population and the cultured strain, respectively. No evidence of a recent bottleneck was detected in the wild population, but it was found in the cultured strain based on the BOTTLENECK test. These primers can be used to understand the demography and to examine genetic differences between the cultured T. albonubes strains and wild populations to help determine conservation and reintroduction strategies.     

Ovarian Hormones Influence Territorial Aggression in Free-Living Female Mountain Spiny Lizards

Females are aggressive in many species but relatively little is known about the hormonal basis of female aggression, especially in free-living animals. Female mountain spiny lizards aggressively defend territories from other females. Previously, we showed that plasma levels of testosterone (T) and estradiol (E) are positively associated with levels of female aggression. Here, we manipulated hormone levels in free-living females and examined aggression expressed by females returned to their natural territories. Females received one of the following: (1) ovariectomy + empty implant (OVEX), (2) ovariectomy + T implant (T-IMP), or (3) sham surgery + empty implant (SHAM). OVEX females had reduced plasma levels of E but not T relative to SHAM females. T-IMP females had elevated plasma levels of T. Levels of display and aggression in OVEX females were reduced relative to SHAM females. T-IMP females had restored levels of display behavior although, unlike SHAM, no T-IMP females expressed the overt aggressive behavior of charging. These data are most consistent with the hypothesis that an ovarian factor such as E promotes female aggression, since ovariectomy reduced both plasma E and aggression but had no effect on plasma T. The results from the T-IMP females are also consistent with this hypothesis if we assume that the effects of T are due to aromatization to E in target tissues. The data do not rule out a role for T in promoting female aggression since T-implants resulted in elevated plasma T and restored display behaviors. This study represents one of the first studies examining the hormonal basis of female aggression in free-living females.     

Polymorphism and division of labour in a socially complex ant: neuromodulation of aggression in the Australian weaver ant,Oecophylla smaragdina

Complex social structure in eusocial insects can involve worker morphological and behavioural differentiation. Neuroanatomical variation may underscore worker division of labour, but the regulatory mechanisms of size-based task specialization in polymorphic species are unknown. The Australian weaver ant, Oecophylla smaragdina, exhibits worker polyphenism: larger major workers aggressively defend arboreal territories, whereas smaller minors nurse brood. Here, we demonstrate that octopamine (OA) modulates worker size-related aggression in O. smaragdina. We found that the brains of majors had significantly higher titres of OA than those of minors and that OA was positively and specifically correlated with the frequency of aggressive responses to non-nestmates, a key component of territorial defence. Pharmacological manipulations that effectively switched OA action in major and minor worker brains reversed levels of aggression characteristic of each worker size class. Results suggest that altering OA action is sufficient to produce differences in aggression characteristic of size-related social roles. Neuromodulators therefore may generate variation in responsiveness to task-related stimuli associated with worker size differentiation and collateral behavioural specializations, a significant component of division of labour in complex social systems.     

Conflict between antipredator and antiparasite behaviour in larval damselflies

 Larval damselflies resist infestation by parasitic larval mites by exhibiting behaviours such as grooming, crawling, swimming, and striking at host-seeking mites. Larval damselflies are known to increase time spent in these behaviours in the presence of mites but reduce time spent in these behaviours in the presence of fish predators. The presence of both fish and larval mites presents an obvious conflict: a larval damselfly may actively avoid parasitism by mites, thus increasing its risk of predation, or it may reduce its activity when fish are present, thus increasing its risk of parasitism. We analysed the behaviour of larval Ischnura verticalis in an experiment where we crossed presence and absence of fish with presence and absence of larval mites. Presence of mites induced a large increase in activity of larval I. verticalis but fish had no effect and there were no interpretable interactions between effects of mites and fish. Subsequent experiments indicated that larval I. verticalis in the presence of both mites and fish were more likely to be attacked and killed by fish than those exposed only to fish. The high activity level of I. verticalis larvae in the presence of both fish and mites may suggest that costs of parasitism are high, or that under field conditions it is rare for larvae to be in the immediate presence of both fish predators and potentially parasitic mites. Received: 28 March 1996 / Accepted: 6 September 1996     

A comparative analysis of aggression in migratory and non-migratory salmonids

This paper represents a comprehensive test of the hypothesis that aggression in salmonids increases with the duration of stream residence. The intraspecific aggression of eleven juvenile salmonids was compared with their normal duration of stream residence. Salmonids maintained in 1 metre tanks and observed over two days could be separated into four groups based on statistical differences between the frequencies of aggressive behaviour. Non-anadromous Salvelinus fontinalis showed the highest levels of aggressive behaviour, followed by Oncorhynchus masou and O. mykiss (rainbow and steelhead trout). Least aggressive were the early migrants O. nerka (kokanee and sockeye salmon) and O. keta. The remaining species, O. rhodurus, O. kisutch, S. leucomaenis pluvius and Salmo trutta formed a moderately aggressive group. Aggressiveness was significantly and positively correlated with the duration of stream residence.     

When supercolonies collide: territorial aggression in an invasive and unicolonial social insect

Some species of ants possess an unusual form of social organization in which aggression among nests is absent. This type of social organization, called unicoloniality, has been studied in only a handful of species and its evolutionary origins remain unclear. To date, no study has examined behavioural and genetic patterns at points of contact between the massive supercolonies that characterize unicoloniality. Since interactions at territory boundaries influence the costs of aggression and the likelihood of gene flow, such data may illuminate how supercolonies are formed and maintained. Here we provide field data on intraspecific territoriality for a widespread and invasive unicolonial social insect, the Argentine ant (Linepithema humile). We observed abrupt and well-defined behavioural boundaries at 16 contact zones between three different pairs of supercolonies. We visited nine of these zones weekly during a six-month period and observed consistent and intense intercolony aggression that resulted in variable, but often large, levels of worker mortality. Microsatellite variation along six transects across territory borders showed that F(ST) values were lower within supercolonies (0.08 +/- 0.01 (mean +/- SE)) than between supercolonies (0.29 +/- 0.01) and that this disparity was especially strong right at territory borders, despite direct and prolonged contact between the supercolonies. Matrix correspondence tests confirmed that levels of aggression and genetic differentiation were significantly correlated, but no relationship existed between geographic distance and either intraspecific aggression or genetic differentiation. Patterns of F(ST) variation indicated high levels of gene flow within supercolonies, but little to no gene flow between them. Overall, these findings are inconsistent with a model of relaxed ecological constraints leading to colony fusion and suggest that environmentally derived cues are not the prime determined of nestmate recognition in field populations of Argentine ants.     

Impact of season and social challenge on testosterone and corticosterone levels in a year-round territorial bird

Plasma testosterone increases during breeding in many male vertebrates and has long been implicated in the promotion of aggressive behaviors relating to territory and mate defense. Males of some species also defend territories outside of the breeding period. For example, the European nuthatch (Sitta europaea) defends an all-purpose territory throughout the year. To contribute to the growing literature regarding the hormonal correlates of non-breeding territoriality, we investigated the seasonal testosterone and corticosterone profile of male (and female) nuthatches and determined how observed hormone patterns relate to expression of territorial aggression. Given that non-breeding territoriality in the nuthatch relates to the reproductive context (i.e., defense of a future breeding site), we predicted that males would exhibit surges in plasma testosterone throughout the year. However, we found that males showed elevated testosterone levels only during breeding. Thus, testosterone of gonadal origin does not appear to be involved in the expression of non-breeding territoriality. Interestingly, territorial behaviors of male nuthatches were stronger in spring than in autumn, suggesting that in year-round territorial species, breeding-related testosterone elevations may upregulate male-male aggression above non-breeding levels. In females, plasma testosterone was largely undetectable. We also examined effects of simulated territorial intrusions (STIs) on testosterone and corticosterone levels of breeding males. We found that STIs did not elicit a testosterone response, but caused a dramatic increase in plasma corticosterone. These data support the hypothesis that corticosterone rather than testosterone may play a role in the support of behavior and/or physiology during acute territorial encounters in single-brooded species.     

Laboratory studies on the effects of thermal change on the behaviour and distribution of juvenile chum salmon in sea water

Schooling chum salmon Oncorhynchus keta were biased towards the water surface (median position <1 m) under isothermal conditions (10° C) in a water column simulator (WCS). Thermal stratification (24/10° C) inhibited upward movement with fish congregating at the thermocline and displaying a clear avoidance of potentially lethal surface waters. A tri-phase model based on piece-wise nonlinear regression was used to describe the distribution shifts of chum salmon during a change from isothermal to thermally stratified conditions. Fish distribution was consistent with thermoregulatory behaviour and exhibited 'attraction', 'preference' and 'avoidance' phases. The thermal preference of 50% of the fish lay between 12·2 and 20·2° C, however, >83·5% of the fish occupied a 'preferred' temperature range of 13·7–17·9° C. The mean temperature at which 50% of chum salmon avoided rising temperature by shifting deeper in the water column and using the cooler thermocline was 20·2° C, and 90% avoidance occurred at 22·9° C. Behavioural responses to thermal stratification were consistent amongst underyearling fish of differing size and age.     

Context matters: female aggression and testosterone in a year-round territorial neotropical songbird (Thryothorus leucotis)

Testosterone promotes aggressive behaviour in male vertebrates during the breeding season, but the importance of testosterone in female aggression remains unclear. Testosterone has both beneficial and detrimental effects on behaviour and physiology, prompting the hypothesis that selection favours an association between aggression and testosterone only in certain contexts in which intense or persistent aggression may be beneficial. We tested this hypothesis in a year-round territorial female buff-breasted wrens (Thryothorus leucotis), by exposing free-living females to experimental intrusions in different social (either single female or male, or paired decoys) and seasonal (pre-breeding or breeding) contexts. Females responded more aggressively to intrusions by females and pairs than to males. However, female intrusions elicited stronger responses during pre-breeding, whereas responses to pair intrusions were more intense during breeding. Territorial females had elevated testosterone levels after female intrusions and intermediate levels after pair intrusions during pre-breeding, but the levels of testosterone remained low after these intrusions during breeding. These results demonstrate seasonal differences in circulating testosterone following territorial aggression in female buff-breasted wrens and are suggestive of differences according to social context as well. Context-dependent elevation of testosterone implies that selection acts directly on female vertebrates to shape patterns of testosterone secretion.     

The effect of dietary docosahexaenoic acid on schooling behaviour and brain development in larval yellowtail

The onset of schooling behaviour and brain growth in larval yellowtail Seriola quinqueradiata larvae fed on Artemia enriched with oleic acid (OA), eicosapentaenoic acid (EPA) or docosahexaenoic acid (DHA) for 10 days was investigated. Larvae from the DHA groups schooled with parallel orientation while those of the EPA groups showed only aggregation. Larvae of the OA groups were dispersed. DHA and EPA groups showed significantly smaller nearest neighbour distance compared with OA groups, and DHA groups showed significantly smaller values of nearest neighbour angle compared with the other groups. The relative volume of the tectum opticus (TO) of the DHA group was significantly larger in two experiments and also the relative volume of the cerebellum (CE) of that group was significantly larger in one experiment. Dietary DHA is probably critically important for the development of the brain, particularly TO, during the larval stages, and that development of TO may be the key factor of the ontogeny of schooling behaviour.     

School Formation and Concurrent Developmental Changes in Carangid Fish with Reference to Dietary Conditions

The ontogeny of schooling behaviour was studied in comparison to the development of sensory and swimming organs and taxis in carangid fish. Striped jack, Pseudocaranx dentex, larvae showed strong phototaxis at 3 days after hatching (3.5 mm in TL) when they developed pigmentation in the retina. Rheotaxis and optokinetic responses were apparent at 4.0-6.0 mm TL as larvae completed development of the basic structure of their eyes. A major inflection of allometric growth occurred at 9 mm, and fin ray formation was completed at these stages. Schooling behaviour, represented by one TL of inter-individual distance and parallel orientation, only appeared at 16 mm TL, and just prior to this behaviour, fish showed mutual attraction through vision at 12 mm TL. Canalization of buccal lateral lines was complete at 18 mm TL, whereas that of trunk lateral lines started at 23 mm TL and was complete at 30 mm TL. With these results, we assumed that critical factors of the ontogeny of schooling behaviour in carangid fish include not only the development of sensory or swimming organs, but also other factors such as development of the central nervous system. To show this, we reared another carangid species, the yellowtail Seriola quinqueradiata, with dietary depletion of DHA (docosahexaenoic acid), which is indispensable for the development of the central nervous system. Although DHA-free fish showed optokinetic response, they did not show schooling behaviour when they attained their schooling size. Tracer experiments using radioisotope labelled DHA showed that DHA is incorporated into the brain, spinal cord, and retina of juvenile fish. Under natural conditions, carangid fish larvae should intake enough DHA through diet to develop schooling behaviour, the fluctuation of dietary quality in zooplankton might therefore influence the development of indispensable antipredatory behaviour. Morphological changes of striped jack occurred in two steps; first at 9-12 mm (fin formation and inflection of allometric growth) and then second at 20-30 mm (scale and lateral line formation), and these changes corresponded with the development of schooling and recruitment to coastal waters, respectively. Since the onset of schooling is the first step of active antipredatory behaviour, we considered that 12 mm TL is the size at which they attain the juvenile period.     

The role of chemoreception in sex recognition by male crickets: Acheta domesticus and Teleogryllus oceanicus

ABSTRACT. Contact chemoreception is important in female recognition by Teleogryllus oceanicus (Le Guillou) males. Antennal contact of female conspecifics, body regions, detached antennae and conditioned substrate elicited mostly courtship responses including courtship songs. Aggressive acts were produced only in response to male conspecifics. Male body regions, detached antennae and conditioned substrate elicited very few courtship or aggressive acts and no songs. This suggests that one or several communication modes, in addition to chemical communication, are necessary to elicit aggressive responses. Acheta domesticus (L.) males cannot rely upon chemical cues for recognition of either sex. Responses to conspecifics suggest that A. domesticus males produce aggressive acts immediately after antennal contact with either sex. Aggressive response to males usually persists, but response to females often switches to courtship. Responses to body regions, detached antennae, and conditioned substrate were few, with courtship and aggressive responses elicited by both male- and female-generated stimuli. The importance of contact chemoreception in cricket communication is suggested by (1) failure of hexane-washed antennae to elicit aggressive or courtship acts, and (2) males spending more time in contact with body regions and conditioned substrates than with corresponding controls. Lack of response to male or female odour-laden air suggests that chemical signals are used by males only if directly contacted. Chemical and other signals supplement the obvious use of acoustic signals for intra- and intersexual communication in these crickets. The importance of multimodal communication in sex recognition is discussed.