Reproductive competition and fecal testosterone in wild male giant pandas (<Emphasis Type="Italic">Ailuropoda melanoleuca</Emphasis>)

The giant panda (Ailuropoda melanoleuca) is notoriously difficult to study in the wild, but its interesting reproductive ecology makes the effort worthwhile. Perhaps more than most species, the panda is energy-limited, which alters the cost/benefit analysis of its reproductive ecology. Using global positioning system/very high frequency radiocollars to locate mating aggregations, we used behavioral observations and fecal testosterone assays to gain insight into male panda reproductive effort and strategies, and test theories relating to reproductive competition. Male pandas initially competed fiercely for access to females that were about to be fertile, but once male competitive status was determined, aggression rates declined. Contact aggression was only observed during the first 2 days of mating aggregations; thereafter, it was replaced with noncontact aggression and avoidance. Agonistic interactions were highly asymmetrical, with contest losers (subordinates) showing less aggression and more avoidance than contest winners (dominants), both before and after contest outcome was established. The competitively superior male displayed mate-guarding tactics and secured all observed copulations. Contrary to theoretical predictions, testosterone levels did not predict aggression levels or contest winners and also were not affected by winning or losing a contest. Body size appeared to be the primary determinant of contest outcome. We discuss our findings in light of theoretical predictions, such as those arising from the “challenge hypothesis,” in the context of the giant panda’s foraging and nutritional ecology.     

Trail discrimination signal of Lasius japonicus (Hymenoptera: Formicidae)

Summary. Trail-following behavior of Lasius japonicus was colony-specific in the field, while trail pheromone activity was not. We found that the footprint substance caused colony-specific trail-following behavior only when working in conjunction with the trail pheromone. The footprint substance alone did not lead the workers to follow trails. The substance consisted mainly of hydrocarbons with composition almost identical to that of cuticular hydrocarbons, except for the absence of n-alkanes. Nestmate workers shared footprint hydrocarbon profiles as well as cuticular hydrocarbons, but the profiles differed among colonies. We therefore consider that the footprint hydrocarbon profiles serve as the trail discrimination signal in L. japonicus.     

Predatory behavior and chemical communication in two Metapone species (Hymenoptera:Formicidae)

Summary. Colonies of two species of Metapone (M. madagascarica, M. new species.) were collected in Madagascar and established in laboratory nests. It could be demonstrated that both species are specialist predators of termites (Cryptotermes kirbyi). During hunting the ants sting the termites and thereby paralyze and preserve the prey alive. In this way prey can be stored in the ant nest for extended periods. During foraging and colony emigrations the ants lay chemical trails with poison gland secretions. Among the seven compounds identified in the venom only methyl pyrrole-2-carboxylate elicits trail following behavior in both Metapone species. Received 11 February 2002, accepted 23 February 2002.     

The honey bee shaking signal: function and design of a modulatory communication signal

This study explores the meaning and functional design of a modulatory communication signal, the honey bee shaking signal, by addressing five questions: (I) who shakes, (II) when do they shake, (III) where do they shake, (IV) how do receivers respond to shaking, and (V) what conditions trigger shaking. Several results confirm the work of Schneider (1987) and Schneider et al. (1986a): (I) most shakers were foragers (at least 83%); (II) shaking exhibited a consistent temporal pattern with bees producing the most signals in the morning (0810–1150 hours) just prior to a peak in waggle dancing activity; and (IV) bees moved faster (by 75%) after receiving a shaking signal. However, this study differs from previous work by providing a long-term, temporal, spatial, and vector analysis of individual shaker behavior. (III) Bees producing shaking signals walked and delivered signals in all areas of the hive, but produced the most shaking signals directly above the waggle dance floor. (IV) Bees responded to the signal by changing their direction of movement. Prior to receiving a signal, bees selected from the waggle dance floor moved, on average, towards the hive exit. After receiving a signal, some bees continued moving towards the exit but others moved directly away from the exit. During equivalent observation periods, non-shaken bees exhibited a strong tendency to move towards the hive exit. (V) Renewed foraging activity after food dearth triggered shaking signals, and, the level of shaking is positively correlated with the duration of food dearth. However, shaking signal levels also increased in the morning before foraging had begun and in the late afternoon after foraging had ceased. This spontaneous afternoon peak has not previously been reported. The shaking signal consequently appears to convey the general message “reallocate labor to different activities” with receiver context specifying a more precise meaning. In the context of foraging, the shaking signal appears to activate (and perhaps deactivate) colony foraging preparations. The generally weak response elicited by modulatory signals such as the shaking signal may result from a high receiver response threshold which allows the receiver to integrate multiple sources of information and which thereby increases the probability that receiver actions will be appropriate to colony needs. Received: 21 March 1997 / Accepted after revision: 30 August 1997     

The behaviour of giant clams (Bivalvia: Cardiidae: Tridacninae)

Giant clams, the largest living bivalves, live in close association with coral reefs throughout the Indo-Pacific. These iconic invertebrates perform numerous important ecological roles as well as serve as flagship species—drawing attention to the ongoing destruction of coral reefs and their associated biodiversity. To date, no review of giant clams has focussed on their behaviour, yet this component of their autecology is critical to their life history and hence conservation. Almost 100 articles published between 1865 and 2014 include behavioural observations, and these have been collated and synthesised into five sections: spawning, locomotion, feeding, anti-predation, and stress responses. Even though the exact cues for spawning in the wild have yet to be elucidated, giant clams appear to display diel and lunar periodicities in reproduction, and for some species, peak breeding seasons have been established. Perhaps surprisingly, giant clams have considerable mobility, ranging from swimming and gliding as larvae to crawling in juveniles and adults. Chemotaxis and geotaxis have been established, but giant clams are not phototactic. At least one species exhibits clumping behaviour, which may enhance physical stabilisation, facilitate reproduction, or provide protection from predators. Giant clams undergo several shifts in their mode of acquiring nutrition; starting with a lecithotrophic and planktotrophic diet as larvae, switching to pedal feeding after metamorphosis followed by the transition to a dual mode of filter feeding and phototrophy once symbiosis with zooxanthellae (Symbiodinium spp.) is established. Because of their shell weight and/or byssal attachment, adult giant clams are unable to escape rapidly from threats using locomotion. Instead, they exhibit a suite of visually mediated anti-predation behaviours that include sudden contraction of the mantle, valve adduction, and squirting of water. Knowledge on the behaviour of giant clams will benefit conservation and restocking efforts and help fine-tune mariculture techniques. Understanding the repertoire of giant clam behaviours will also facilitate the prediction of threshold levels for sustainable exploitation as well as recovery rates of depleted clam populations.     

Sex attractant of an arctiid moth (Utetheisa ornatrix): A pulsed chemical signal

Summary The sex attractant pheromone produced by the female of the moth Utetheisa ornatrix was shown to contain Z, Z, Z-3, 6, 9-heneicosatriene. The compound, whose structure was confirmed by synthesis, proved active in electroantennogram and field bioassays. Pheromone emission occurs discontinuously, in the form of short pulses (pulse repetition rate=1.5±0.2 pulses/s). It is argued that such temporal patterning — which had not previously been demonstrated for an airborne chemical signal — can provide close-range orientation cues to the male moth as it seeks out the female.     

The neglected scent: on the marking function of urine in Lemur catta

In ring-tailed lemurs (Lemur catta) urine deposition can be combined with two different tail configurations: (i) tail held up in an evident display (urinate tail-up, UT-up); (ii) tail slightly raised to avoid its impregnation with urine (urinate tail-down, UT-down). We used both signaller- and receiver-based approaches to search for functional differences between these two kinds of urine deposition. We predicted that UT-up might be a complex signal combining olfactory and visual cues. We carried out observations and scent tests on four captive groups of ring-tailed lemurs. Group members sniffed/licked UT-up scents more frequently than UT-down ones. Moreover, UT-up showed peak levels during the mating season whereas UT-down did not. These findings suggest that urine can play a role in intra-group reproductive communication. Lemurs more frequently performed UT-up in a few drops and UT-down in streams. Recognition experiments clearly showed that individuals can discriminate between urine of their own group and urine from a foreign group (a necessary prerequisite for the use of urine in inter-group communication). The possible function of UT-up in inter-troop communication was supported by the higher frequency of this pattern along a fence separating two of the study groups. Moreover, in the presence of a dummy, the frequency of UT-up increased significantly. In conclusion, UT-up is a complex signal with multiple characteristics. By using different sensory channels, UT-up provides different types of information (location and signaller quality) and contains multiple messages directed both at group-members and neighbouring groups....to conceal this letter, the Minister had resorted to the comprehensive and sagacious expedient of not attempting to conceal it at all (from the Purloined letter E. A. Poe, 1845)     

The unsung songs of great tits (Parus major): learning neighbours' songs for discrimination

Summary Many territorial birds are capable of recognising neighbours from their songs. In this paper we investigate the timing of learning of neighbours' songs for recognition. This is the first time that the timing of discrimination learning has been studied explicitly by experiment in the field. We show that discrimination learning is not limited to early life and contrast this with the timing of song learning for performance. We also show that the ability to recognise new neighbours is inversely related to the number and similarity of neighbour's songs which have previously been experienced. This suggests pro-active memory interference and provides the first field evidence for memory constraints on discrimination.     

The chemical basis for trail recognition in Lasius nipponensis (Hymenoptera: Formicidae)

Summary. In this paper we report on the chemical basis for trail recognition in Lasius nipponensis. On and near trails in the field, workers became aggressive against conspecific intruders and succeeded in protective contests, while intruders usually avoided confrontation. Such asymmetric interactions were also observed in the laboratory when two non-nestmate workers were sequentially placed in a clean glass dish, while mutual aggression was observed when they were placed simultaneously. Asymmetric aggression was also observed when the workers were placed in a dish previously conditioned by other workers, or when the dish was treated with the hexane rinse of the conditioned dish. The rinse contained a series of hydrocarbons, in which components and proportion were almost identical to those of the cuticular hydrocarbons of the workers, except for a lack of n-alkanes. Amounts of the hydrocarbons that remained on the dish were very small when all the tarsi of the workers were incapacitated. We therefore consider that the hydrocarbons are not adhesion of the cuticular hydrocarbons but secretion from the tarsi of workers, and serve as a key signal for trail recognition in this ant.     

Nestmate discrimination in social wasps: The role of exposure to nest and nestmates (Polistes fuscatus,Hymenoptera: Vespidae)

Summary In the laboratory, gynes (potential queens) of the paper wasp, Polistes fuscatus, exposed to both their natal nest and female nestmates (presumably their sisters) discriminate between female nestmates and unrelated females. However, gynes not exposed to their natal nest or conspecifics and gynes exposed only to female nestmates do not discriminate between female nestmates and unrelated females. Thus, the presence of the nest appears to be a requisite for the development of nestmate discrimination.     

Sperm motility and longevity in the giant cuttlefish, Sepia apama (Mollusca: Cephalopoda)

The sperm kinetics and fertilisation literature in marine invertebrates is heavily biased toward free-spawning species. Nonetheless, many species (e.g. cephalopods) transfer and/or fertilise gametes in confined external spaces or internally, creating very different selective pressures on sperm storage, sperm longevity and hence sperm competition. Here we report the results of an investigation into the effects of sperm age, water temperature and sperm concentration on sperm motility in the giant cuttlefish (Sepia apama). Significant positive correlations were found between percent motility and sperm concentration, and between sperm motile speed and sperm concentration. Mean percent motility of cuttlefish sperm suspension was still 9% eight hours after being released from the spermatophore and diluted into filtered seawater at 12°C (ambient field temperature during the spawning season). Sperm resuspended from spermatangia taken from (mated) females in the field were motile for up to 100 hours. When spermatophores were stored at 4°C motility was still observed in resuspended sperm after two months. Our results show that spermatangia and spermatophores can retain and release live sperm for long periods. The observed longevity of sperm in S. apama greatly increases the potential for sperm competition in this species.     

Consequences of communal gall occupation and a test for kin discrimination in the aphid Tamalia coweni (Cockerell) (Homoptera: Aphididae)

This study investigated the consequences of communal gall induction on individual and group fitness in the aphid Tamaliacoweni. The possibilities that kin discrimination and foundress density are factors favoring communal gall occupation were examined. Clonally produced aphid foundresses were collected to create two treatments: clonal groups and groups of less closely related individuals. These were confined on suitable host plant tissue to compare their respective propensities towards communal behavior. There were no significant differences in the frequencies of communal gall occupation; therefore, active kin discrimination by T.coweni foundresses apparently does not play a role in their communal behavior, within the context of this experiment. In a second experiment, aphid foundress density on the host plant was manipulated in three treatments and was correlated with the frequency of communal gall occupation. Individual fitness was inversely related to the mean number of foundresses per gall. These results suggest that communal gall occupation does not necessarily represent mutual cooperation but may instead be the outcome of competition for limited gall sites on the host plant. However, natural selection at the clonal level may favor communal gall occupation under the conditions of resource limitation and high within-group relatedness. Received: 26 September 1997 / Accepted after revision: 28 March 1998     

Regulation of worker reproduction in bumblebees (Bombus terrestris): workers eavesdrop on a queen signal

In the annual bumblebee Bombus terrestris, the onset of queen-worker conflict over male production is seasonally and socially constrained. Workers will do better if they start to reproduce (the so-called competition phase) only after ascertaining that larvae are committed to gyne development but before the season ends because they gain more by rearing sister-gynes than their own sons. Here, we tested two nonmutually exclusive hypotheses as to what triggers the onset of worker reproduction: Workers can directly monitor larval development and/or workers eavesdrop on the queen signal that directs gyne development. Exposing workers to gyne larvae through a double mesh did not advance the competition phase compared to control colonies. However, when workers, but not the queen, were allowed contact with gyne larvae, both the competition phase and gyne production were advanced. Thus, while larvae do not emit a volatile pheromone that discloses their developmental route, the physical contact of workers with such larvae triggers early competition phase. However, workers exclusively exposed to worker larvae (colonies prevented from producing gyne larvae) started to reproduce at the same time as control colonies. Replacing the resident queen with an older queen (from gyne-rearing colonies) advanced the competition phase, irrespective of worker age. The results are consistent with the hypothesis that workers eavesdrop on the queen pheromones. This is adaptive because it allows workers a broader time-window for reproduction and thus to gain fitness from rearing both sister-gynes and sons before the season ends without affecting colony development.     

Effects of feeding history on mobilisation and deposition of body constituents and on growth in juvenile Ambassis vachelli (Pisces: Chandidae)

Young Ambassis vachelli (Richardson) 40 to 50 d old, were used in a laboratory experiment to test the effects of starvation and subsequent re-feeding on body constituents and growth. Fish in three laboratory treatments (fed continuously; fed for 9 d and then starved for 15 d; starved for 9 d and fed for 15 d), were compared to fish from a local field population. Different body stores were mobilised at different times and rates during starvation. Carbohydrates were mobilised from the onset of starvation and were depleted after 3 d. Lipid and protein were mobilised at an increasing rate from the onset of starvation. The mortality in starved fish was relatively high (up to 70%) until re-feeding. Upon feeding, all body stores were restored rapidly in fish that were starved, with carbohydrate levels displaying an overshoot (carbohydrate level exceeding normal levels) in comparison to the levels in continuously fed fish. After 15 d of feeding, the starved fish had levels of constituents similar to those in continuously fed fish. Fish that were fed and subsequently starved were able to maintain themselves for at least 15 d prior to death, indicating a better degree of starvation resistance than fish without a history of feeding. This implies that feeding history in the early life of a fish is important in growth and survival but that young fish may have growth regimes flexible enough to survive relatively long periods of starvation.     

Studies on the defensive behaviour ofHypselodoris species (Gastropoda: Nudibranchia): Ultrastructure and chemical analysis of mantle dermal formations (MDFs)

The mantle dermal formations (MDFs) ofHypselodoris webbi (D'orbigny, 1839), andH. cantabrica (Bouchet and Ortea, 1980) are globular sub-epidermal structures located in the cephalic and caudal regions. Histologically they consist of an accumulation of vacuolate cells surrounded by a basal lamina and an outer muscular capsule. Chemical analysis ofH. webbi MDFs reveals the presence of high concentrations of longifolin, a well known deterrent furanosesquiterpenoid that had been previously isolated from this species. In the present paper it is demonstrated that the great majority of longifolin accumulated in the mantle ofH. webbi is stored in the MDF vacuolar cells. This finding strongly suggests that such structures act as chemical weapons against predation, mainly protecting vital organs such as the head, rhinophores and gills.     

The effect of temperature-salinity combinations on survival and growth of juvenile Patiriella pseudoexigua (Echinodermata: Asteroidea)

Adult Patiriella pseudoexigua were collected in October 1989 from Wanlitung, Taiwan and then induced to spawn in the laboratory. Post-metamorphosed juvenile P. pseudoexigua were reared on a diet of benthic algae Navicula sp. at 25°C and salinity (34). Six weeks after metamorphosis, juvenile P. pseudoexigua at ca. 400 m in radius were reared on a diet of benthic algae Navicula sp. at different combinations of temperatures (20, 25, 30°C) and salinities (26, 30, 34) for 40 d. Both temperature and salinity had a significant effect on juvenile survival and growth. Juveniles survived best (>90%) at 25°C and 34 and grew best (to ca. 750 m in radius) at 30°C and 34. Variation in juvenile size was small immediately after metamorphosis and increased with time.     

The effect of temperature and thermal distribution on glycolysis in two rockfish species (Sebastes)

Studies on the effects of temperature on the activities of Embden-Meyerhof (EM) glycolysis, and the hexose monophosphate shunt (HMP) in fishes have dealt mainly with exotic and/or acclimated fishes. This study reports the effects of short-term reductions in temperature on EM and HMP activity in two closely related species of temperate fishes (Sebastes spp.) and its possible relation to the thermal distribution of the species. Thermal distribution data were collected by SCUBA for S. mystinus and S. serranoides in King Harbor, Redondo Beach, California, USA. Activities of the pathways were determined in liver-tissue studies, using glucose-¹⁴C and liquid scintillation techniques following the method of Hochachka (1968) with modifications. The data were analyzed by distribution-free methods. Tissue studies indicated HMP activity in both species at lowered temperature (5°C), but only in S. serranoides at 15°C. Results indicate that S. mystinus is capable of instantaneous temperature compensation, possibly related to its tendency to occupy cold waters.