A Comparative Study of Agonistic Behaviour in Hairy-footed Gerbils (Genus Gerbillurus)

Intraspecific agonistic behaviour was investigated by means of staged encounters in Gerbillurus paeba paeba, G. p. exilis, G. tytonis, G. setzeri and G. vallinus. Four groups of behaviour were identified by motivational analysis: i.e. “exploratory and solitary”, “aggressive”, “submissive”, and “sexual” behaviour. Frequencies of acts and levels of interaction differed among species and sexes. Cluster analysis of behaviour data revealed three clusters which agree partially with the karyology and taxonomy of the genus. G. p. paeba and G. tytonis were very aggressive and highly active; female G. p. exilis and G. setzeri less so; while male G. p. exilis and G. vallinus were least aggressive and active, performing more contact-promoting behaviour than other species. A range of social types is exhibited from solitary (G. p. paeba and G. tytonis) through semi-tolerant (G. setzeri and female G. p. exilis) to tolerant (male G. p. exilis and G. vallinus).     

A Comparative Study of the Behaviour of Six Taxa of Male and Female Gerbils (Rodentia) in Intra- and Interspecific Encounters

Frequencies and sequences of visually-identifiable acts were investigated in male-female combinations of gerbils of the taxa Tatera brantsii, Gerbillurus paeba paeba, G. p. exilis, G. tytonis, G. setzeri, and G. vallinus. Sexual dimorphism of behaviour was most apparent in T. brantsii, G. setzeri, and G. vallinus, in which following, anogenital sniffing, and mounting were significantly more frequently performed by males than females. Females of all taxa presented and darted, while males did not perform these acts. Analysis of first-order transitions of acts in intraspecific encounters revealed remarkable similarity among species in the sequences identified as most significantly deviating from expected frequency. Species-specific patterns of behaviour were detected by discriminant analysis. G. p. paeba, G. p. exilis and G. vallinus were not clearly distinguished by discriminant analysis, and presented a behaviour pattern characterized by low levels of aggression and high levels of huddling behaviour. G. tytonis, G. setzeri and T. brantsii were characterized by low levels of huddling, G. tytonis performed the largest number of upright postures of all taxa, G. setzeri watched more than other taxa, and T. brantsii nasal sniffed less frequently than Gerbillurus taxa. Differences among taxa were more apparent in males than in females. Interspecific encounters were staged between G. p. paeba and G. p. exilis, G. p. paeba and G. tytonis, G. setzeri and G. vallinus. Males showed more evidence of changed behaviour in interspecific encounters than females. More sexual and less agonistic behaviour was detected in intraspecific encounters than in interspecific encounters, but this was statistically significant in only 5 of 12 combinations. Visual, tactile and olfactory communication during physical contact with another individual may complement auditory communication in promoting conspecific recognition in southern African gerbils.     

Ultrasonic Vocalizations of Six Taxa of Southern African Gerbils (Rodentia: Gerbillinae)

Ultrasonic calling during male-female encounters between individuals of the same species was investigated in six taxa of southern African gerbils, namely Tatera brantsii, Gerbillurus paeba paeba, G. p. cxilis, G. tytonis, G. setzeri, and G. vallinus. Vocalizations were detected by means of a bat detector utilizing a superheterodyne signal converter and a countdown circuit. Signals were recorded at audible frequencies and analysed with a sonograph. All taxa vocalized at ultrasonic frequencies by means of strongly modulated frequency “sweep” calls, which differed among taxa in duration, maximum and minimum frequency. “Clicks” were emitted by G. p. paeba and G. p. exilis, and G. tytonis emitted a “stutter” vocalization which consisted of a series of clicks. Long modulated “whistles” were identified from G. vallinus and T. brantsii at lower frequencies than “sweep” calls. Only one call type, a “sweep” call which differed in duration and frequency from all other taxa, was identified in G. setzeri. Cluster analysis was applied to the data using 7 acoustic characters. G. p. paeba and G. p. exilis displayed the highest similarity level between taxa and differed only in frequency of “sweep” vocalizations. G. paeba, G. tytonis and G. setzeri formed one cluster, while G. vallinus and T. brantsii formed a separate cluster. Numbers of calls in interspecific encounters were non-significantly less than in intraspecific encounters in all taxa except G. p. paeba, in which more vocalizations were recorded in inter- than intraspecific encounters. It is not clear whether species discrimination, measured by numbers of vocalizations in interspecific encounters, occurs.     

Behaviour Associated with Ultrasonic Vocalizations in Six Gerbilline Rodent Species

Encounters were staged between male and oestrus-induced females of the same or different taxa of the species/subspecies Gerbillurus paeba paeba, G. p. exilis, G. tytonis, G. setzeri, G. vallinus and Tatera brantsii. Each encounter lasted 20 min; converted ultrasonic emissions were recorded on videotape simultaneously with visual images. Videotapes were transcribed and sequences of acts and vocalizations were recorded. The frequency of acts was compared separately with the number of ultrasonic vocalizations during and after acts by χ² contingency table analysis, followed by partitioned χ² to identify individual acts which contributed significantly to the overall χ² value. Vocalizations occurred significantly more frequently than expected during sexual and/or huddling behaviour in all species-groups; and significantly less frequently than expected during exploratory, watching and/or submissive behaviour. Vocalizations were uncommon during investigative behaviour in all taxa, and aggressive behaviour in Gerbillurus taxa, but occurred significantly more frequently than expected after these behaviour categories in all taxa. Analysis of vocalizations immediately following acts verified the interpretations of function of vocalizations during acts. Ultrasonic vocalizations clearly play an important role in the communication of six taxa of southern African gerbils.     

Capture-release field observations of free-ranging chacma baboons,Papio ursinus,focusing on the behaviour of the sub-adult male segment of the baboon social unit

An adult female and three age-graded male chacma baboons,Papio ursinus, were captured and then artificially introduced into a recipient troop of free-ranging baboons of the same species. Two of the transplanted individuals, the youngest juvenile male and the sub-adult male, were accepted into the troop while the oldest juvenile male and the adult female were aggressively chased off by the sub-adult and adult males of the troop, and they subsequently returned more than 10 km to their home troop. Acceptance of the first two individuals was dependent on the gradual reduction of aggressive behaviour of the sub-adult males. This suggested that successful troop transfers in the wild, occurring more frequently than traditionally believed, may be at least partially contingent on the behaviour of the sub-adult male segment of the baboon social unit. Supported by a Graduate Research Grant from the University of Cape Town.     

Body Size and the Maintenance of Reproductive Isolation in Stickleback,Genus Gasterosteus

The blackspotted stickleback Gasterosteus wheatlandi and the widely studied threespine stickleback G. aculeatus are sympatric throughout the former’s range and share many aspects of life history and reproductive behaviour. These two species differ significantly in size, with G. wheatlandi of both sexes measured at approximately 60% of the standard length of their G. aculeatus counterparts. This study concentrated on G. wheatlandi courtship behaviour and investigated its role in the maintenance of reproductive isolation with G. aculeatus. Specifically, the roles that (1) female body size plays in influencing male courtship preferences and (2) male body size and behaviour play in female courtship preferences were investigated through dummy and live conspecific and heterospecific stimulus presentations. Male G. wheatlandi courtship preferences are consistent with previously described patterns for G. aculeatus. Males of both species preferentially approach and court the larger of two simultaneously presented live or dummy females. Thus, the smaller G. wheatlandi males are indiscriminate with respect to assortative mate choice; not only preferring to approach and court more fecund conspecific females but, more significantly, G. aculeatus‐sized females. In contrast, females of both species demonstrate strong assortative courtship preferences. When presented with pairs of flask‐enclosed males, females of both species preferentially orient and court the conspecific male over the heterospecific. Similarly, when presented with a conspecific male and a heterospecific male presented singly, females prefer to enter the nest of the conspecific. Systematic analysis of the interactions between these pairs of fish (one male, one female) demonstrates that the breakdown of courtship in heterospecific courtship occurs late in the courtship sequence when the widely differing forms of male leading behaviour results in drastically differing female responses. I suggest that, as previously described in G. aculeatus, the supernormality effect plays a significant role in mediating adaptive mate choice behaviour in G. wheatlandi. However, the added element of a larger sympatric species introduces a possible cost in time and energy devoted to courting heterospecific, and sympatric, females that the larger G. aculeatus do not likely incur. There is substantial evidence from many sympatric G. aculeatus species pairs that there is assortative mate choice based on size and/or courtship behaviour. Courtship trials suggest a more pervasive role for females in assortative mate choice. Whether it is male body size per se, or in combination with behaviour, morphology or other cues, is unresolved in the present study.     

Male Courtship Pheromones Affect Female Behaviour in the Swordtail Characin (Corynopoma riisei)

Pheromones constitute an important cue used by both males and females during courtship. Here, we investigate the effect of male pheromones on female behaviour in the swordtail characin (Corynopoma riisei), a species of fish where males have a caudal pheromone gland which has been suggested to affect female behaviour during courtship. We subjected female C. riisei to male courtship pheromones and investigated the effect on both female behaviour and brain serotonergic activity levels compared to a control group. While no difference in serotonergic activity was found, the pheromone‐treated females showed lower stress levels compared to the control group. Furthermore, pheromone‐treated females increased locomotor activity over time, while a decrease in locomotor activity was observed in the control group. These results suggest that the male courtship pheromones may serve to reduce female stress and increase female activity, possibly to aid males in gaining access to females and facilitating sperm transfer.     

Maternal behaviour and neonatal development in three species of Namib Desert rodents

Gerhillurus paeba paeba, G. tytonis and Pelromyscus collinus inhabit arid areas of southern Africa. Several litters of each species were bred in the laboratory; aspects of maternal behaviour and development of neonates are reported. Pelromyscus collinus has a smaller litter size and the young are less altricial at birth than young of G. paeba or G. tytonis. Gerhillurus tytonis young have the slowest rate of physical and behavioural development, while P. collinus young develop most rapidly. Young of P. collinus nipple-cling from birth, but those of G. paeba and G. tytonis do not. This dichotomy may be related to phylogeny, mode of locomotion, and/or nesting habits.     

The effects of male harassment on mating duration in the seed bug,Togo hemipterus

Harassment on mating pairs by solitary males is usually considered an attempt by the male to (1) take over the female, (2) guard the female against further insemination (when the solitary male has previously copulated with this female), or (3) influence mating duration. Paired males of a seed bug repel harassment on mating pairs by solitary males by firmly grasping females using their legs and/or genital claspers; in this way, mating duration is prolonged. Male fertilization success increases as mating duration increases. Males of the seed bug, Togo hemipterus (Scott) (Heteroptera: Lygaeidae), use seminal substances to inhibit female remating. These substances induce protracted female refractory periods and are transferred to the females in a time‐dependent manner. Consequently, mating duration has important effects on fitness in this species. We observed harassment on T. hemipterus mating pairs by solitary males, and examined conflicts between paired and solitary males over mating duration. None of the solitary males were able to take over a mating female, and this may be due to the unique male genital structure in this species. All conflicts over mating duration resulted in wins by the paired males over the solitary males. Paired males prolonged mating durations, whereas severe harassment on mating pairs by solitary males shortened durations. We show that even though there is no immediate reward for the solitary male (i.e., it is unable to take over the mating female), this harassment behavior may be adaptive.     

Reproductive parasitism: male and female responses to conspecific and heterospecific intrusions at spawning in a mouth‐brooding cichlid Ophthalmotilapia ventralis

A rare form of alternative reproductive behaviour without simultaneous parasitic spawning was observed in Ophthalmotilapia ventralis, a lekking mouth‐brooding cichlid from Lake Tanganyika. Floater males attempted to sneak opportunistically into the territory to actively court the female, while the owner (bourgeois male) defended the territory against other potential intruders. Floater males had more body fat than territory owners and generally higher condition factors. In field experiments, the response of bourgeois males and courted females was tested towards floaters and egg predators (a catfish Synodontis multipunctatus) present in the territories. Territory owners responded aggressively particularly to floaters, and female responsiveness to bourgeois male courtship tended to decline when floaters were present. The potential influence of reproductive parasitism on sexual selection in mouth‐brooding cichlids is discussed.     

Information on Resource Quality Mediates Aggression between Male Madagascar Hissing Cockroaches, Gromphadorhina portentosa (Dictyoptera: Blaberidae)

Male Madagascar hissing cockroaches, Gromphadorhina portentosa Schaum (Dictyoptera: Blaberidae) have a well‐defined dominance hierarchy that has been assumed to explain the outcome of most competitive interactions. We studied whether males of this species would alter their level of aggression towards unfamiliar rivals as a function of changing resource availability and value – two factors that are key to aggression levels in non‐hierarchical species. We quantified male aggression as three variables (aggressive state – behaviours measured by their duration; aggressive act – behaviours measured by their frequency of occurrence; aggressive latency – the latency to first aggressive behaviour, either state or act) and tested for any context‐specific variation within each by manipulating both territorial status (males were either residents or intruders) and access to mates (female present or absent). Both the presence of a female and territorial status affected male aggression towards rivals as measured by duration of aggressive state. Highest levels of aggression were displayed by residents when a female was present. These results show that inter‐male aggression in G. portentosa is tuned to the immediate expected payoff from fighting, and not exclusively aimed at establishing dominance relationships (which can affect future payoffs).     

Sex-biased aggression and male-only care at sea in Brünnich's Guillemots Uria lomvia and Razorbills Alca torda

In several groups in the order Charadriformes, biparental care is followed by a period of male‐only care. Several hypotheses attempting to explain extended male parental care in shorebirds do not fit the Alcini. In a previous study of Brünnich’s Guillemots Uria lomvia and Razorbills Alca torda, we did not find support for female‐biased parental effort at the breeding site that would lead to males being in better condition to care for chicks at sea. However, in both species, males spent more off‐duty time at the breeding site than females, suggesting greater involvement in the defence of egg or chick, breeding site and mate. We predicted that there would be a male bias in size and aggressive behaviour associated with parental roles. To test this, body size and aggression of attending male and female Brünnich’s Guillemots and Razorbills were measured during incubation and brooding on the Gannet Islands, Labrador. Parental aggression was measured using natural observations of all agonistic interactions and, in Razorbills only, in situ responses to presentations of a predator model. In both species, males were significantly larger than females in culmen and gape length. Guillemot males initiated agonistic interactions more frequently than females during incubation. In contrast, female Guillemots were subjected to aggression more frequently than males and as a result were involved in more fights. In addition, the few chicks that were seen to die were being attended by single females. During the brooding period, Razorbill males responded aggressively to intruders more frequently than females, made more aggressive responses than females, and responded aggressively more frequently and more intensely than females to a predator model. In both species there was a similar male bias in morphology and behaviour that is consistent with male parents being more capable of protecting their chick, a probable advantage to chick survival during the uniparental care phase of some Charadriformes.     

Factors affecting aggressive behaviour of spawning migratory males towards mature male parr in masu salmon Oncorhynchus masou

This study examined whether dominant migratory males (adopting fighter tactics) of the masu salmon Oncorhynchus masou would more aggressively attack large mature male parr (adopting sneaker tactics) as large mature male parr are expected to have the potential to cause a greater decrease in fertilization success. The frequency of aggressive behaviour was not related to the body size of males, and it increased with the frequency of interactions with mature male parr. The fertilization success of mature male parr was much lower than migratory males, and no relationship was observed between fertilization success and aggressive behaviour. The low fertilization success of mature male parr, despite infrequent aggressive behaviour by migratory males, indicates that there might be little benefit for migratory males to attack mature male parr more aggressively according to their body size.     

Influence of Adult Size on Mate Choice in the Solitary and Gregarious Parasitoids, Cotesia marginiventris and Cotesia flavipes

Female and male mate choice in relation to adult size were examined for the solitary and gregarious parasitoids, Cotesia marginiventris (Cresson) and Cotesia flavipes Cameron, respectively. In addition, male precopulatory behaviors were observed for evidence of male competition or a large-male advantage in mate acquisition. Male parasitoids are not known to offer female mates direct benefits, but females that mate high quality males may obtain indirect benefits, such as offspring that are more successful in obtaining mates. Female choice experiments for C. marginiventris found that large males approached females first more frequently than small males, and that females mated large males significantly more often than small males. Male choice experiments for C. marginiventris did not demonstrate a male preference for female size. In contrast, female choice experiments with C. flavipes found that females mated equally with large or small males, while male choice experiments showed that males attempted copulation and mated more frequently with smaller females. Male competition was not observed in the gregarious species C. flavipes, but competition in this gregarious parasitoid could be moderated by dispersal.     

Differences in size and energy content affect the territorial status and mating success of a neotropical dragonfly

In male odonates, both size and fat content are related to territory defence and mating success. Males that are larger and have higher energy reserves win relatively more disputes for territory and attract more females. Wing colour has also been regarded as a mechanism that influences agonistic behaviour between males, as wing pigmentation might be regarded as a sign of male quality. In this study, we analysed whether a set of male physical (body size and wing colour), physiological (body fat content) and behavioural (disputes between males) characteristics were involved in the territory defence and mating behaviour of the neotropical dragonfly Zenithoptera lanei Santos, 1941 (Anisoptera: Libellulidae). Males were characterised as territorial whenever they warded‐off other males and remained in the same place within the pond for two consecutive days. In general, these territorial males were larger and had more abdominal and thoracic fat, engaged in pursuits more frequently, spent more time on sexual behaviour and female guarding, and mated more in comparison to subordinate males. By evaluating whether the percentage of wing area covered by black ink influenced male behaviour, we found that territorial males tended to act aggressively towards other males whose wings were partially painted, and sexually towards females irrespective of wing area painted. In Z. lanei, both body size and fat content play a role in defining territoriality. By subduing competitors and dominating preferred locations within high‐quality sites, these males are likely to be visited by females and engage in mating.     

Solitary and social bees as pollinators of Wahlenbergia (Campanulaceae): single-visit effectiveness, overnight sheltering and responses to flower colour

Solitary bees often form specialised mutualisms with particular plant species, while honeybees are considered to be relatively opportunistic foragers. Thus, it may be expected that solitary bees are more effective pollinators than honeybees when foraging on the same floral resource. To test this, we studied two Wahlenbergia species (Campanulaceae) in South Africa that are visited by both social honeybees and solitary bees, and which are shown here to be genetically self-incompatible and thus reliant on pollinator visits for seed production. Contrary to expectation, the solitary bee Lipotriches sp. (Halictidae) and social bee Apis mellifera (Apidae), which were the two most frequent visitors to flowers of the study species, were equally effective pollinators in terms of the consequences of single visits for fruit and seed set. Both bee species preferentially visited female phase flowers, which contain more nectar than male phase flowers. Male solitary bees of several genera frequently shelter overnight in flowers of both Wahlenbergia species, but temporal exclusion experiments showed that this behaviour makes little contribution to either seed production or pollen dispersal (estimated using a dye particle analogue). Manipulation of flower colour using a sunscreen that removed UV reflectance strongly reduced visits by both bee groups, while neither group responded to Wahlenbergia floral odour cues in choice tests. This study indicates that while flowers of Wahlenbergia cuspidata and W. krebsii are pollinated exclusively by bees, they are not under strong selection to specialise for pollination by any particular group of bees.     

Female response to predation risk alters conspecific male behaviour during pre‐copulatory mate guarding

Mating behaviour often increases predation risk, but the vulnerability within mating pairs differs between the sexes. Such a sex difference is expected to lead to differences in responses to predation risk between the sexes. In the two‐spotted spider mite Tetranychus urticae, males engage in pre‐copulatory mate guarding because only the first mating results in fertilisation. We investigated (i) whether pre‐copulatory pairs are more conspicuous to the predatory mite Phytoseiulus persimilis than solitary females, (ii) whether the vulnerability to the predator differs between sexes within the pre‐copulatory pair, (iii) whether each sex of T. urticae responds to predation risk during pre‐copulatory mate guarding and (iv) whether T. urticae's response to predation risk affects predator behaviour. Because T. urticae females are immobile during pre‐copulatory mate guarding, we observed male behaviour to evaluate effects of predation risk. We found that the predators detect more pre‐copulatory pairs than solitary females and that more females than males of the pre‐copulatory pairs are preyed upon by the predators. The preference of spider mite males for pre‐copulatory pairs versus solitary females was affected by whether or not the female had been exposed to predators during development. Male T. urticae exposed to predation risk did not alter their behaviour. These results suggest that only the most vulnerable sex, that is the female, responds to predation risk, which modifies male behaviour. Regardless of T. urticae females’ experience, however, P. persimilis detected more T. urticae pre‐copulatory pairs than solitary females, suggesting that pre‐copulatory mate guarding itself is dangerous for T. urticae females when these predators are present. We discuss our results in the context of sex‐dependent differences in predation risk.     

Predator‐inspection behaviour in female three‐spined sticklebacks Gasterosteus aculeatus is associated with status of gravidity

Differences in predator‐inspection behaviour between gravid and non‐gravid female as well as between male and female three‐spined sticklebacks Gasterosteus aculeatus were investigated. Gravid females confronted with a live rainbow trout Oncorhynchus mykiss showed bolder inspection behaviour than non‐gravid ones. The behaviour of gravid females was comparable with that of males, maybe because both face a high risk of predation. The results indicate that antipredator behaviour in female G. aculeatus is not fixed but adjusted to their reproductive state.     

Female receptivity and male mate-guarding in the jewel spiderGasteracantha minax thorell (Araneidae)

In field populations, several male jewel spidersGasteracantha minax Thorell (Araneidae) may be found at the periphery of the orb web of a female, indicating that males may compete for fertilisation success. Laboratory experiments revealed that virgin femaleG. minax readily remate shortly after their first mating. However, they appear to enter a refractory period between 1 and 24 h postmating and respond aggressively to courting males. Males that have mated with a female initially defend her from rival males but cease to do so after the onset of the refractory period. These data can be interpreted within the context of mate-guarding and sperm competition. There is marked size dimorphism in this species, which may be the result of selection for protandry. In contrast with other orb-weaving spiders, this selection pressure does not seem to be counterbalanced by selection for larger male size through either sexual cannibalism or male-male competition.     

Male Aggression,Female Mimicry and Female Choice in the Rove Beetle,Aleochara curtula (Coleoptera,Staphylinidae)

Causes, consequences and pheromonal regulation of male contest and female choice in the staphylinid beetle, Aleochara curtula (Goeze), have been investigated in the field and in the laboratory. At the feeding and mating site (carcass) the sex ratio is male biased. Polyandry is affected by prolonged copulations, spermatophore plugs and anti-aphrodisiac pheromones transferred from the male, and by female repulse behaviour as well. Aggression of competing males leads to expulsion of inferior males from the carcass. Young, starved and multiply mated males, which need access to the food resource, produce the female sex pheromone. They release homosexual responses, but also avoid intrasexual aggression. On the other hand, females behave aggressively towards individuals bearing the female sex pheromone or repulse their copulatory attempts. Those males of insufficient physiological condition produce a lighter spermatophore and fertilize less eggs. The adaptive significance of female mimicry, male mating tactics, and female choice is discussed.