Female Mate Preference for Males Having Long and Symmetric Fins in the Bower-holding Cichlid Cyathopharynx furcifer

Female mate preference in a bower-holding cichlid, Cyathopharynx furcifer, was studied in Lake Tanganyika. Most males held territories with crater-shaped bowers in sand, but some males held territories without bowers. Territories were distributed adjacently and females visited them to spawn. After engaging in circling behaviour with the male, a female deposited eggs in the bower. Soon after spawning, the female picked the eggs up into her mouth and brooded them in places away from male territories. Female mate choice appeared to follow three steps: 1) females visited only bower-holding male territories, and more frequently visited territories of males that performed courtship displays at a higher frequency and had longer pelvic fins; 2) females preferred to start circling with males having longer and more symmetrical pelvic fins; 3) females chose males with more symmetrical pelvic fins as their mates. Less than 7% of females that visited male territories spawned eggs in the bowers. In contrast to other bower-holding species, bower size did not correlate with male reproductive success in C. furcifer. Bowers may therefore be essential as spawning sites or may function as a species recognition character for females. Female choice may be dependent instead on males having long and symmetrical pelvic fins apparent during the circling behaviour carried out in the bowers.     

Preliminary study on female choice of mates versus sites in a wrasse, Cirrhilabrus temminckii

Female choice of mates versus sites was studied in a wrasse, Cirrhilabrus temminckii. Males had territories within a restricted area on a rocky slope at which females visited and pair-spawned pelagic gametes. Females visited several males or territories before spawning, suggesting the opportunity of female choice. Of the four characteristics of territorial males examined—body size, ratio of pelvic fin length to body size, courtship, frequency and territory depth—only territory depth was significantly correlated with daily mating success of males. The former three male characters were not related to territory depth. These results suggest that female C. temminckii chooses deep sites rather than specific mates in mating.     

Female preference for males that have exclusively marked or invaded territories depends on male presence and its identity in the subterranean rodent Ctenomys talarum

Territorial scent‐marking provides chemical records of male competitive interactions that are available to females, who gain valuable information to assess and identify best quality partners. In this context, the solitary subterranean rodent tuco‐tuco (Ctenomys talarum) offers excellent possibilities to evaluate the effects of male exclusive scent‐marking of territories on female assessment. For evaluation, we used wild caught individuals of C. talarum, manipulated their scent marks within the territories in captive conditions and staged preference tests where females were able to choose between exclusive and invaded territories. The evaluation was performed in two scenarios considering the identity of the intruder scent mark: territories invaded by a strange male and territories invaded by a neighbour male. Females investigated the chemical cues deposited on the substrate of the exclusively marked territory more frequently. Next, females displayed equal interest to scent samples of both males presented in a Y‐maze. Finally, when females could gain access to both individually isolated males and their scent‐marked territories, they spent more time within invaded territories despite they visited them with the same frequency. Moreover, females tried to get in contact by scratching the mesh of the owner of the invaded territory more frequently. We found that females of C. talarum evaluate the homogeneity (exclusiveness) of scent marks within a male territory and then show preferences in relation to the identity of the intruder's scent –whether strange or neighbour.     

The Effect of Variation in Male Sexually Dimorphic Traits on Female Behaviour in Pronghorn (Antilocapra americana)

Several sexually dimorphic morphological, behavioural, and territory characteristics of male pronghorn, Antilocapra americana, are highly variable. To test the hypothesis that females respond to variation among males for mate choice, I examined the relationship between these male characteristics and female congregation size and variability. Increased movement up to and during rut and differential female congregation on the territories of males with favoured characteristics were detected as predicted. Individual females increased their home ranges nearly fourfold from summer to rut and visited 88% of the territories in the study area. Female congregation size was related to male phenotype but not to territory characteristics. A canonical correlation analysis revealed that not only were several male traits simultaneously related to female congregation size and variability, but the importance of these traits varied across years. However, male courtship, cheek-patch size, and scent marking behaviours were consistently correlated to female congregation patterns across years. Female congregation was significantly related to male courtship, but there was no correlation between former territory owner's and newcomer's attraction of females. This finding is consistent with the hypothesis that females were primarily assessing males instead of territories.     

Why is Female Choice not Unanimous? Insights from Costly Mate Sampling in Marine Iguanas

Females do not unanimously choose the single 'best' male, even when female choice is strong, such as in leks, or in polygynous mating situations. A possible explanation is that females base their choices on limited information, perhaps because gathering information is costly. We tested this hypothesis by continuously observing individual female marine iguanas throughout the mating period in order to document the information they gathered about each potential mate. Females actively visited approximately five additional males during the 3 d prior to copulation, compared to the males seen on their normal foraging routes. Females were more likely to visit large-bodied males, but preferentially copulated with the male that had the highest display rate of all males they visited. Females that mated on a dense territory cluster mated with more active males than did those that mated on dispersed territories. However, females on a dense cluster also lost more body mass, potentially as a consequence of high rates of interaction with males. This mass loss may represent an important cost and result from postural changes in response to male attention. Such costs may explain why females only gather a certain amount of information and why females on dispersed territories choose less active mates. Lack of complete information introduces subjectivity into female choice: what is perceived as best by one female may not be perceived as best by another. Thus, lack of complete information may prevent unanimity of female choice.     

Supplementation of male pheromone on rock substrates attracts female rock lizards to the territories of males: a field experiment

Background

Many animals produce elaborated sexual signals to attract mates, among them are common chemical sexual signals (pheromones) with an attracting function. Lizards produce chemical secretions for scent marking that may have a role in sexual selection. In the laboratory, female rock lizards (Iberolacerta cyreni) prefer the scent of males with more ergosterol in their femoral secretions. However, it is not known whether the scent-marks of male rock lizards may actually attract females to male territories in the field.

Methodology/Principal Findings

In the field, we added ergosterol to rocks inside the territories of male lizards, and found that this manipulation resulted in increased relative densities of females in these territories. Furthermore, a higher number of females were observed associated to males in manipulated plots, which probably increased mating opportunities for males in these areas.

Conclusions/Significance

These and previous laboratory results suggest that female rock lizards may select to settle in home ranges based on the characteristics of scent-marks from conspecific males. Therefore, male rock lizards might attract more females and obtain more matings by increasing the proportion of ergosterol when scent-marking their territories. However, previous studies suggest that the allocation of ergosterol to secretions may be costly and only high quality males could afford it, thus, allowing the evolution of scent-marks as an honest sexual display.     

Nectar and flower production of Lobelia telekii inflorescences, and their influence on territorial behaviour of the scarlet-tufted malachite sunbird (Nectarinia johnstoni)

The alpine zone of Mount Kenya is a typical equatorial, high-altitude habitat with a harsh environment, large fluctuations in physical conditions and an impoverished flora and fauna. A common flowering plant is Lobelia telekii , which has large inflorescences with up to 2000 flowers. The long hanging bracts of this plant ameliorate the physical environment around the flowers, damping the fluctuations in temperature and relative humidity. The scarlet-tufted malachite sunbird ( Nectarinia johnstoni ) is a highly active nectarivore that obtains most of its food from the nectar of L. telekii. Sunbirds are more likely to visit the younger, male flowers at the apex of the inflorescence, which are larger and contain twice as much sugar as the older, female flowers lower down the spike. Male flowers will gain from multiple visits by exporting more pollen, while female flowers probably need to be visited only a few times for successful fertilization. Male sunbirds were resident on their territories all year and vigorously defended them from conspecifics. All territories contained about four times the number of flowers visited by the territory occupants in a day. Males on territories with a large number of flowers suffered more intrusions than males on low quality territories and spent more time in flight. After an intrusion the resident male frequently fed near where the intrusion took place. Males that subsequently attracted mates defended about twice as many flowers as males that did not breed, although undefended inflorescences were present. Some males apparently defended territories suitable for a pair, others only for a single bird.     

Male territoriality and mating system in the European beewolf Philanthus triangulum F. (Hymenoptera: Crabronidae): evidence for a “hotspot” lek polygyny

The evolution of the mating system of a species is strongly influenced by the spatial and temporal distribution of females and/or resources. Here, we describe aspects of the territorial behavior of males of a solitary digger wasp, the European beewolf (Philanthus triangulum) and characterize the mating system. We show that beewolf males establish small territories that do not contain any resources essential to females. These territories are intensively scent-marked with a pheromone from a cephalic gland and are defended against intruders in combat flights. We provide evidence that scent-marking constitutes a chemical display and that the pheromone serves to attract receptive females and, thus, represents a sex pheromone. Using spatial statistics, we show that beewolf territories are clumped in space both with respect to other male territories and, more importantly, with respect to female nesting sites. Additionally, the proportion of days a territory is occupied by a male is correlated with the number of female nests in the vicinity. Taking into account that beewolf males do not defend or provide resources essential to females, but merely display chemically to attract females for mating in an aggregation of territories close to female nesting sites, we conclude that the European beewolf exhibits a hotspot lek polygyny with female nesting sites constituting “hotspots” for lek formation.     

The social organization of feral peafowl

Peafowl are usually reported to have a mating system based on harem defence by adult males. In a small feral population near Oxford, males defended small (<1 ha) territories while females remained in one flock that ignored male territory boundaries. After mating, females become solitary. At no time did a female associate selectively with one male or remain within his territory, nor did males attempt to follow or guard female groups. Two out of four males were seen to mate. These differed from the other two in being neither very old nor very young; they held territories smaller than that of the young male and were no larger or longer-tailed. However, they spent more time displaying. We suggest that peafowl have a mating system similar to a lek: males defending small, clumped territories visited by females for mating.     

Female mate choice in the carpenter bee Xylocopa varipuncta (Hymenoptera: Anthophoridae)

Females of the carpenter bee Xylocopa (Neoxylocopa) varipuncta Patton exhibit highly active mate choice. During the spring in central Arizona, males of this species hover at shrubs and trees on or near ridgelines in late afternoons. Occasionally, females fly to and closely approach pheromone-releasing males on their territories, but in 65 of 85 approaches the female left without copulating. Some females visited several males in a few minutes. Copulation only occurred when the female landed on a spot that the resident male rubbed with his body upon the female's arrival. Males did not attempt to disrupt courtship or copulation by other males. In 1988 one creosote bush proved especially attractive to males, as measured by the frequency with which the site was occupied simultaneously by more than one male. The majority of close approaches by females occurred in this one bush. Simultaneous occupation of hovering sites was very rare in 1989. In this year, there was a weak but significant correlation between the frequency with which different sites were visited by intruder males and by females. These results support the hypothesis that males are able to identify locations most likely to attract potential mates.     

Tactic for bower acquisition by male cichlids,Cyathopharynx furcifer,in Lake Tanganyika

A tactic for bower acquisition by male cichlids,Cyathopharynx furcifer, was studied in Lake Tanganyika. Males held crater-shaped bowers of sand within their territories, which were located adjacent to each other. Females visited bowers to spawn eggs, subsequently brooding the eggs in their mouths. Some individuals held territories without bowers near those of males with bowers. Although only the males with bowers succeeded in reproduction, they were apparently exhausted by bower maintenance and reproduction, and often deserted their bowers. These were soon occupied by males which had held nearby territories without bowers, such males then engaging in reproductive activity. Although the construction of new bowers by males was observed, such was unsuccessful, probably because too much energy was required and/or the optimal space for bowers was limited. Direct aggression by males without bowers against bower-holding males to deprive the bowers was not observed, possibly due to the high costs of such competition. Therefore, waiting for bower desertion by other males may be the best bower acquisition tactic for males without bowers.     

Chemosensory discrimination of male age by female Psammodromus algirus lizards based on femoral secretions and feces

Chemical communication plays an essential role in several social and reproductive behaviors of many animals. In lizards, the main sources of semiochemicals are femoral or pre‐anal gland secretions and feces. In male lizards Psammodromus algirus, there are age‐related differences in the chemical composition of femoral gland secretions and in the reproductive strategies, with older males defending territories and females, while younger males adopting sneak‐mating strategies. Females flee more often from mating advances of young males than from those of old males, which are more successful in obtaining matings. This suggests that age discrimination of males may be important for females. We tested here whether females showed differential chemosensory responses to chemical cues (femoral gland secretion and feces) of males of two age classes, and whether females use information from substrate scent marks of males of different ages to select where to stay. We found that females elicited more tongue‐flicks to the secretion and feces of old males than to control or secretion and feces of young males. Also, the time spent by females on a scented paper depended on the treatment, suggesting that females tended to spend more time on scent marks made with femoral secretions of old males. Adult females seemed capable to discriminate between young and old males based on chemical cues alone and showed more interest in scents of old males. However, substrate scent marks did not seem to entirely determine site selection by females, suggesting that females might need additional cues to perform the choice. These results can be explained by the different age‐dependent reproductive strategies of males, which can affect differentially to females.     

Social organization and spawning in the Atlantic sharpnose puffer,Canthigaster rostrata (Tetraodontidae)

Synopsis Social organization and spawning in the sharpnose pufferCanthigaster rostrataere studied on a reef in the San Blas Islands, Panama. Sexes were dimorphic. In mixed coral and rubble habitat, females defended territories against other females and small males. From one to six female territories were included within the territories of certain large males. These haremic males visited their females and patrolled their territories throughout the day. Smaller, non-haremic males occupied territories or home ranges within or adjacent to those of haremic males or were wanderers. Spawning between a haremic male and a territorial female occurred within the female's territory. The female prepared an algal nest into which demersal eggs were deposited. There was no parental care. Eggs were spherical, translucent, and measured approximately 0.66 mm in diameter. Larvae were about 1.4 mm TL and closely resembled those of other species ofCanthigaster.     

Scent marking in a territorial African antelope: I. The maintenance of borders between male oribi

Scent marking is ubiquitous among the dwarf antelope and gazelles of Africa, but its function has been the subject of debate. This study examined preorbital gland scent marking in the oribi, Ourebia ourebi, a territorial African antelope. Several hypotheses for the function of scent marking by territorial antelope were tested with observational data. Of these, the hypotheses that scent marking is driven by intrasexual competition between neighbouring males, and that marks serve as an honest advertisement of a male's ability to defend his territory from rivals, were supported best. Thirty-three territorial male oribi on 23 territories marked most at borders shared with other territorial males, and territorial males marked more often at borders shared with multimale groups than at borders shared with a single male. This suggests that males perceived neighbouring male groups as a greater threat to territory ownership than neighbouring males that defended their territories without the aid of adult subordinates. Marking rate was unrelated to territory size or the number of females on adjacent territories, but males with many male neighbours marked at higher rates than those with fewer male neighbours. These results suggest that the presence of male neighbours has a greater effect on the scent marking behaviour of territorial antelope than has been considered previously. Copyright 1999 The Association for the Study of Animal Behaviour.     

Reproductive Behaviour of Japanese Tubesnout, Aulichthys japonicus (Gasterosteiformes), in the Natural Habitat Compared with Relatives

We observed significant sexual dimorphism in size (females > males) and in colour in Japanese tubesnout, Aulichthys japonicus, in Aburatsubo Bay, central Japan, and Otsuchi Bay, northern Japan. This species is known to spawn its eggs into ascidians. We observed reproductive behaviour of tubesnout in the natural environment in both bays. All the males established territories at harbours or rocky shores and showed conspicuous nuptial colouration with metallic green or blue on their snout and bright yellow on the body. Females had a dark brown body with remarkable white spots. The male territories varied somewhat in location and extent on a daily basis. The individual male size and territory size were not correlated. The females usually stayed in the seaweed beds where no males were present. Males and females were distributed separately, and the gravid females visited the male territories in groups to spawn. The territorial males courted the gravid females by rushing and returning, and tried to lead the females to the particular place of their territories by tail beating. We discuss nuptial colouration, courtship behaviour and territoriality in comparison to the related species; Aulorhynchus flavidus, Hypoptychus dybowskii, and gasterosteid fishes.     

Temporal and Spatial Distribution of Male Scent Marks in the Polygynous Greater Sac-Winged Bat

Scent marks are relatively long-lived signals that can be perceived by conspecifics when the producer is absent. Therefore, it is often not obvious to whom the signal is directed. In daytime roosts of the polygynous greater sac-winged bat, males scent mark territories with facial gland secretions. Territories are a valuable resource for males, as they offer exclusive courtship opportunities, which results in increased male reproductive success and, consequently, increased male–male competition over territories. The information encoded in male scent marks could, therefore, be either directed at females as part of an olfactory courtship display or at male competitors as part of territorial behaviour. We expected territorial males to scent mark in the morning, shortly before females return to the territory and close to female roosting sites, if scent marks are directed at females as part of the courtship display. And we expected harem males to scent mark at the territory boundaries, where male–male encounters are most likely to occur, if scent marks are directed at male competitors. We found that males marked more frequently in the afternoon, at a time when all females have already left the territory, and harem males marked at the territory boundaries and not inside their territory in the area where females roost. At boundaries males fan volatiles from specialised wing sacs towards competitors outside the territory. Scent marking of male Saccopteryx bilineata might therefore be congruent with the assessment-hypothesis, which states that scent marks offer intruders the possibility to make an olfactory assessment of the territory owner without direct physical interaction. Thus, scent marks of male S. bilineata are most likely influenced by male–male competition and not by female choice.     

Female mate choice and mate search tactics in a sex role reversed population of the peacock blenny Salaria pavo (Risso, 1810)

Behavioural observations were carried out on a sex role reversed population of the blenny Salaria pavo to investigate possible mate choice behaviour of the females. Females mated with relatively larger males, with a larger head crest, anal gland and genital papilla, which had more eggs in their nests and that courted the female. Thus, these male traits were potentially assessed by females when choosing their mates. In order to mate, females visited one to four males (eight females spawned with the first male they had visited and eight visited more than one male). The majority of the females spawned with the last male visited but three of them returned to mate with males visited previously (two with the penultimate visited male). Additionally, the outcome of the first visit of each female depended on the quality of the males: males that were accepted on this first visit had larger head crest than rejected males. Therefore, the mate searching model that best fitted the data was a threshold tactic that allowed the searching individual to return to any potential mate visited before, i.e. that allowed for complete recall. Moreover, three females returned to mate with males rejected earlier during the search, which indicates that they lowered their thresholds. These results suggest that females use a 'one step decision' tactic to search for mates. In this tactic, females mate with males that satisfy an adjustable threshold criterion, balancing the quality of the mates expected to find in the next step of the search and the effort of finding them.     

The Behavioral Ecology of Libellula luctuosa (Burmeister) (Anisoptera: Libellulidae): I. Temporal Changes in the Population Density and the Effects on Male Territorial Behavior

A single population of a common pond dragonfly, Libellula luctuosa, was studied at a site where the density of males increased dramatically during the breeding season. Early in the summer one active male was found on each territory on the pond. Satellite males were only occasionally found on the territories. Later in the season the number of males per territory increased so that two or more males simultaneously defended on many of the territories, and several satellite males occupied each of the territories. The number and rate of female visitations per day did not change over the summer. These factors resulted in a change in the operational sex ratio with variations in male density. Male behavior was also altered with increasing population density. As male density increased, males were less likely to be seen perching on their territories and more likely to be seen performing aggressive acts such as chasing nearby territorial males and chasing intruders. At high male density, the duration of territorial behaviors was shorter than at low male density. Thus, the percent of a time budget spent in any one activity did not change despite the change in number of males present. Male activity in L. luctuosa is not strictly determined by the opportunity for aggression. Costs of aggression associated with territoriality are minimized by maintaining flexible territorial behaviors.     

(1S,3S,7R)-3-methyl-alpha-himachalene from the male sandfly Lutzomyia longipalpis (Diptera: Psychodidae) induces neurophysiological responses and attracts both males and females

Lutzomyia longipalpis adult males form leks on or near hosts and release (1S,3S,7R)-3-methyl-alpha-himachalene from their tergal glands to lure females to the same site for mating and feeding. Here we have examined whether the male-produced attractant could also serve as a male aggregation stimulus. High resolution chiral capillary gas chromatography analysis of male tergal gland extracts, synthetic (1S,3S,7R)-3-methyl-alpha-himachalene, and a synthetic mixture of all isomers of 3-methyl-alpha-himachalene, was coupled to electrophysiological recordings from ascoid sensillum receptor cells in antennae of male and female sandflies. Receptor cells of both sexes responded only to the main component of the male tergal gland extract that eluted at the same retention time as (1S,3S,7R)-3-methyl-alpha-himachalene. Furthermore, of the eight 3-methyl-alpha-himachalene isomers in the synthetic mixture only the fraction containing (1S,3S,7R)-3-methyl-alpha-himachalene, co-eluting with an isomer of (1S*,3S*,7S*)-3-methyl-alpha-himachalene, elicited an electrophysiological response from male and female ascoid sensillum receptor cells. Both males and females flew upwind in a wind tunnel towards a filter paper disk treated with either 4-6 male equivalents of the tergal gland extract, pure (1S,3S,7R)-3-methyl-alpha-himachalene or the synthetic mixture of eight isomers. This indicates that (1S,3S,7R)-3-methyl-alpha-himachalene derived from L. longipalpis males may have a dual function in causing male aggregation as well as serving as a sex pheromone for females.     

Spawning behavior and haremic mating system in the corallivorous butterflyfish,Chaetodon trifascialis, at Kuroshima Island, Okinawa

Territorial and spawning behavior ofChaetodon trifascialis were investigated on a small patch of reef at Kuroshima Island, Okinawa, Japan. Three males and 8 females inhabited the reef, each individual defending a territory against conspecifics of the same sex. Each male territory included 2 or 3 female territories. In the daytime, each male frequently visited the females living in its territory. At dusk in the full or new moon periods, courtship began within the female territories, pair spawning subsequently occurring within or near those territories. When a male actively courted a female in the territory of a second male, the latter male immediately chased off the intruder. Thus, mating occurred only between a male and females living in former's territory. This is the first report of a haremic mating system among butterflyfishes (Chaetodontidae).