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.     

Coexistence of permanently territorial cichlids of the genus Petrochromis through male-mating attack

Coexistence mechanisms of permanently territorial fishes proposed hitherto have been mainly based on interspecific competition. To test the hypothesis of the coexistence through male-mating aggression (Kohda, 1995a), spacing patterns of feeding territories of three coexisting herbivorous cichlids, Petrochromis polyodon, P. trewavasae and P. famula, were investigated in a homogeneous habitat at a shallow rocky shore in Lake Tanganyika. All three species maintained individual feeding territories that were defended against both conspecifics and congenerics and rarely overlapped either intra- or interspecifically. Territories of three species were scattered and covered almost all rocky bottoms. Territorial attacks usually occurred near the border of feeding territories. But male P. polyodon, the largest and socially most dominant fish, attacked conspecific males far beyond their feeding territories, which were widely separated. Removal of some territory owners suggested that such attacks result in the separate distribution of male P. polyodon territories. The interstitial space between P. polyodon feeding territories resulting from their mating aggression can be occupied by subordinate congeneric fishes. Male-mating attack of dominant males of Petrochromis will facilitate the coexistence of other congeners.     

Male dimorphism, territoriality and mating success in the tropical damselfly, Paraphlebia zoe Selys (Odonata: Megapodagrionidae)

The tropical damselfly Paraphlebia zoe has two male morphs: a black-winged (BW) male which is associated with territorial defense of oviposition sites; and a hyaline-winged (HW) male similar in appearance to females, and, compared to the black morph, less frequently found defending territories. In a wild population of this species, we first assessed the relationship between phenotypic traits [male morph, size and territorial status (being territorial or non-territorial)], their role on mating success, and the degree to which a particular territory may contribute to male mating success. Second, to relate a physiological basis of being territorial we compared both morphs in terms of muscular fat reserves and thoracic muscle, two key traits related to territory defense ability. Males of both morphs defended territories although the BW males were more commonly found doing this. BW males were larger than HW males and size predicted being territorial but only within HW males (territorial males were larger) but not in BW males. Male mating success was related to territorial status (territorial males achieved a higher mating success), but not to morph or size. Furthermore, territory identity also explained mating success with some territories producing more matings than others. The BW morph stored more fat reserves which may explain why this morph was more likely to secure and defend a place than the HW morph. However, the HW morph showed higher relative muscle mass which we have interpreted as a flexible strategy to enable males to defend a territory. These results are distant to what has been found in another male dimorphic damselfly, Mnais pruinosa, where the advantage of the non-territorial morph relies on its longevity to compensate in mating benefits compared to the territorial morph.     

The Haremic Mating System and Mate Choice in the Wide-Eyed Flounder, Bothus podas

The social structure and reproductive behaviour of the wide-eyed flounder, Bothus podas, was studied in the coastal waters around the Azorean Islands. Both sexes are territorial throughout the year. Adult males defend large territories, which include several smaller female territories. Intraspecific agonistic behaviour was frequent and differed between sexes: males were more aggressive towards other males, while females were only aggressive towards each other and juveniles. During the reproductive season and only at dawn, territorial males court and mate successively with females in their territories, and females seem to show mating fidelity to their dominant male. Such territoriality and mating patterns indicate a haremic social system in the wide-eyed flounder. In order to identify potential factors influencing female mate choice acting on this haremic system, we examined male mating success and some of its potential correlates. We found no evidence for female preference for any of the males' physical or territory characteristics. However, courtship effort was strongly correlated with the total number of attempted and successful spawnings, indicating that females seem to mate preferentially with males that court them more vigorously. Thus, our data suggest that courtship plays an important role in determining male mating success in the wide-eyed flounder and, that it may possibly serve as an honest indicator of male `quality' for female choice.     

Mating tactics and male wing dimorphism in the damselfly,Mnais pruinosa costalis selys (Odonata: Calopterygidae)

Males of the damselfly,Mnais pruinosa costalis, exhibit wing color dimorphism: one form has orange wings, and the other hyaline wings which resemble female wings. The former is usually territorial and the latter uses sneaky mate securing tactics. When orange-winged males failed to establish territory, they became floaters that day. Hyaline-winged males perched around their territories and often, formed in tandem without any apparent courtship behavior when they found females. Their copulation frequency was higher and copulation duration longer than those of territorial males. A few females oviposited without remating. Total oviposition duration of females with which a hyaline-winged male mated was more than 32 min per male on average in a day Females that copulated with hyaline-winged males often remated with orange-winged residents before oviposition. Total duration of oviposition bouts of females after mating with floaters was short (15 min), while that with territorial residents was long (66 min). As a result, total oviposition duration of females with which an orange-winged male mated was about 40 min in a day. The reproductive success of the hyaline-winged males may be similar to that of the orange-winged males.     

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.     

Territoriality and nutritional condition in Cynotilapia afra (Gunther) and Pseudotropheus zebra (Boulenger), cichlidae in Lake Malawi National Park, Malawi

The standard length, diurnal activities, territory sizes, and areas over which individuals foraged and the nutritional condition of territorial and non-territorial Cynotilapia afra and Pseudotropheus zebra were compared. Results show that territorial tenureship in these fishes does not depend on the male size, implying that aggressiveness, experience and motivation are more important in the maintenance of territory. However, terri-toriality in C. afra (t = 10.93, P < 0.05) and P. zebra (t = 3.31, P < 0.05) manifests itself in the reduction of nutritional condition because territorial males engage in energetically more demanding activities, e.g. courtship displays, fertilizing ripe females and chasing intruders that trespass into their territories. Territorial C. afra (t = 4.77; P < 0.05) and P. zebra (t = 5.89; P < 0.05) also fed over significantly smaller areas and spent significantly less time feeding than did non-territorial males. The biological significance of territoriality in these fish species is therefore not food intake, but mate attraction and reproductive success for which they trade off their nutritional condition. However, there might be a nutritional threshold below which the cost out-weighs the benefit. Hence, territorial males in poorer health abandon their territories in order to regain their condition.     

The effects of male body size on territorial and mating success in the landmark-defending fly Hermetia comstocki (Stratiomyidae)

Abstract.

• 1 Males of Hermetia comstocki Williston compete for territorial control of certain agaves and yuccas. Winners copulate with females that visit these plants solely to acquire a mate.
• 2 Males vary in body weight by more than an order of magnitude and larger flies almost always defeat smaller ones in aerial contests for control of landmark territories.
• 3 The mean body size (as measured by wing-length) was significantly greater for males retaining residency at a site for at least one hour compared to males unable to do so. Likewise, males able to return to a perch site in the study area on more than one day were larger on average than males unable to do so.
• 4 Male preferences for landmark territories remained similar across years. Large males dominated the perch landmarks most likely to be occupied by males and most likely to be visited by females.
• 5 Despite the fighting and territorial advantages enjoyed by large males, the mean size of males found mating with females was not significantly larger than that of the general population.
• 6 The apparent failure of large males to secure a statistically significant mating advantage may be a statistical consequence of the small sample size of males observed mating. On the other hand, any mating advantage of large males may be reduced because (a) receptive females visit many different landmarks, (b) females mate with the first male they encounter at a landmark, regardless of his size, (c) there are usually many vacant landmarks available for smaller males, and (d) even popular territories are often open to small males, thanks to the low site-tenacity of territory owners.

     

Influences on variation in territorial tenures of male white-faced dragonflies (Leucorrhinia intacta) (Odonata: Libellulidae)

Some individuals in species with extended periods of territorial occupancy may change territory locations within a single bout of territorial activity. Length of occupancy of mating territories among males in a local population of white-faced dragonflies (Leucorrhinia intacta) varied from more than 6 h to 15 min or less. Males with short tenures often established territories in several locations on the pond during a day. Several hypotheses have been proposed to explain shifting territorial sites rather than remaining in a single site during one bout of territoriality. We attempted to test the hypothesis that males shift to leave low-quality sites. Site quality may be affected by costs of defense in relation to intruder rate and the mating benefits of holding the territory. To test whether variation in these possible effects of benefits and costs of territoriality influenced tenure, we manipulated local quality of oviposition substrate and perch density. The quality of oviposition substrate, but not perch density, influenced both potential benefits and costs of territoriality. Female density was higher in areas with good substrate, but so were rates of males intruding into the territories, rates of chasing by territorial males, and local density of territorial males. More matings occurred in areas with good substrate, but among males with tenures of 15 min or more, mating success per male and tenure lengths did not differ statistically among treatments. Defense costs were low for all treatments and perhaps were not an important influence on tenure duration. Territorial males in this population probably adjusted local density to expected mating success by initial choice of site rather than by varying tenure length. Variation in tenure length at a site resulted, in part, from stochastic external factors, such as predation attempts.     

The role of male territories in the mating system of the Yellow-shouldered Widowbird Euplectes macrourus

The territorial system and breeding biology of the Yellow-shouldered Widowbird Euplectes macrourus (Ploceidae) was investigated in western Kenya. Yellow-shouldered Widowbirds had a resource-defence polygynous mating system: males defended large (mean = 0.95 ha) territories and built the coarse framing for the nests in tall grass. Males had up to five females nesting per territory. Females provided nearly all parental care except for a territorial male seen feeding a fledgling: the first observation of paternal care in the wild for this genus. There was considerable variation in territory size, but the cause of this variation remains unknown: territory size was not related to potential indicators of territory quality, such as grass height and abundance, did not relate to male morphology (mass, size and ornament size) or territorial behaviour (boundary displays and singing) and did not affect female preferences. Although resources (territories and nests) were defended by the males, observations that males frequently fed outside their territories and formed communal roosts during the breeding season suggest that this species represents a transitional stage between typical resource-defence polygyny and lek breeding.     

Competition for landmark territories among male Polistes canadensis (L.) (Hymenoptera: Vespidae): large-size advantage and alternative mate-acquisition tactics

Alternative male mating tactics of insects at landmarks (leks)have only rarely been investigated. Some males of the paperwasp, Polistes canadensis (L.), were territorial at small treesalong the crests of dry ridges in Santa Rosa National Park,Costa Rica. Territories did not contain nests or resources forwhich females foraged. Contrary to other "hilltopping" species,male P. canadensis competed most intensely for territories insaddles along these ridges rather than at the highest points.Nonterritorial males patrolled small areas of the ridge line,following a path that took them to a number of territories.Many males switched between territoriality and patrolling, suggestingthat both size-related tactics belong to one conditional strategy.Males that were territorial on 2 or more days were larger thanthose that were territorial on only 1 day, and these in turnwere larger than permanent patrollers. Moreover, the mean sizeof territorial males was positively correlated with two measuresof territory attractiveness, suggesting that larger males monopolizepreferred sites. Mean age of territorial males was also relatedto territory attractiveness, but males of intermediate age claimedthe most attractive territories.     

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.     

Androgen control of social status in males of a wild population of stoplight parrotfish, Sparisoma viride (Scaridae)

In the protogynous stoplight parrotfish (Sparisoma viride), large males defend territories that encompass the home-ranges of several mature females. However, high-quality habitat is in short supply, such that smaller, competitively inferior males do not defend territories. We investigated the role of 11-ketotestosterone (11KT) and testosterone (T) in the regulation of territorial behavior in a wild population of a protogynous reef fish, the stoplight parrotfish, at Glover's Reef, Belize. Radioimmunoassay of plasma samples from individuals of known social status revealed that nonterritorial males have lower levels of T and 11KT than territorial males. Nonterritorial males allowed access to vacant territories underwent pronounced increases in T and 11KT. When sampled 1 week after territory acquisition, levels of T and 11KT in new territorial males were significantly higher than the levels in established territorial males, but by 3 weeks after territory acquisition, there was no significant difference. We further investigated the hypothesis that such short-term increases in androgen levels are a response to intense male-male interactions during territory establishment. Simulated territorial intrusion promoted increased plasma levels of both T and 11KT while access to vacant territories without neighboring territorial males did not. These findings suggest that the endocrine system plays a role in fine-tuning the levels of territorial aggression exhibited by male stoplight parrotfish. We discuss these results in light of recent theory in behavioral endocrinology.     

Male spawning success and female choice in the mottled triplefin,Forsterygion varium (Pisces: Tripterygiidae)

A detailed analysis was made of the variation in spawning success among male Forsterygion (э Tripterygion) varium. The aim was to assess whether females were mating selectively and, if so, to examine the basis of that choice. Eggs were laid on small rocks within the males' territories. Females did not distribute eggs randomly among males. Some males guarded as many as 23000 eggs in one breeding season, while others experienced little or no breeding success. Male spawning success was positively correlated with both their size and some characteristics of their territories, namely the amount of spawning substratum and the presence of a large boulder adjacent to the territory. Experimental manipulations indicated that female choice of males was based on these features. Addition of small rocks into the territories of previously unsuccessful males resulted in a significant increase in spawning success. Conversely, spawning success of the most successful males decreased when small rocks were removed from their territories. When successful males were removed, each territory was occupied by small males; however, there was no significant change in the spawning success of these males. This experiment suggested that, although territory quality was important, female mating decisions could also be influenced by male characteristics such as body size.     

A comparative study of territoriality in intertidal and subtidal blennioids (Teleostei, Blennioidei)

This paper presents a comparative analysis of territoriality in three intertidal (Lipophrys pholis, Coryphoblennius galerita, Salaria pavo) and two subtidal (Tripterygion delaisi, Parablennius pilicornis) blennioid fishes. Focal-animal observations of males guarding eggs showed that: (i) intertidal species had smaller territories that were less frequently patrolled; (ii) in subtidal species feeding was limited to the territory, while in intertidal fishes a substantial proportion of the feeding acts occurred outside the defended area; (iii) intertidal species spent less time out of the nest and showed a lower level of locomotory activity; (iv) subtidal species were subjected to a higher number of territorial intrusions with more species intruding their territory, some of them potential egg predators; (v) subtidal species performed several water column displays that played a significant role in courtship, while in intertidal fishes these displays are almost absent. Signalling without the presence of a female was almost absent in intertidal species. It is argued that intertidal fishes minimize the time spent out of the nest and the loss of contact with the substrate, features that may be of high survival value in conditions of strong turbulence. Although intertidal species have a reduced time available for feeding, this may be compensated by lower levels of locomotory activity, territorial defense and risk of egg predation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Short-term changes in plasma levels of hormones during establishment and defense of a breeding territory in male song sparrows, Melospiza melodia

When territorial male song sparrows are captured and removed from their territories, previously unmated and nonterritorial males will take over those vacant territories within 12-72 hr. Plasma levels of testosterone are elevated in these replacement males as well as in their neighbors. Since the latter already have territories, it is suggested that the agonistic interactions over territory boundaries, or behavioral stimuli from challenging males, rather than ownership of a territory per se, stimulates secretion of testosterone. To test this hypothesis further, male song sparrows were challenged by experimental simulation of a territorial intrusion. This procedure involved placing a caged male song sparrow in the center of the subject's territory and playing tape recorded conspecific song through an adjacent speaker. Responding males were then captured at intervals after onset of the intrusion. Plasma levels of testosterone were significantly higher in males exposed to experimental territorial intrusion than they were in controls, supporting the hypothesis that behavioral stimuli emanating from an intruding male can act as supplementary information stimulating secretion of testosterone.     

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.     

Microsatellite determination of male reproductive success in a natural population of the territorial ornate dragon lizard, Ctenophorus ornatus

It is now evident that the genetic mating system can be very different to the observed mating system. However, it is less well known what makes particular individuals more (or less) successful than expected from the observed system. In this study the observed territorial structure of a field population of the agamid lizard, Ctenophorus ornatus, was compared with the mating system as evidenced by microsatellite parentage assignment. This study also investigated whether any male traits predicted reproductive success. Sixty-five per cent of clutches were sired at least partially by a male other than the main territory-holding male and 35% of clutches were sired by a male with no overlap of the female's territory. Multiple paternity was moderately frequent at 25% of clutches. Male chest patch size predicted territory size and the number of females in the territory, but did not predict reproductive success. Instead, male head depth and body size were independently related to the number of offspring sired. As male head depth also predicted the number of females in a territory, these males are likely to be gaining increased reproductive success as a consequence of the higher number of females in their territories. Larger body size males, however, did not have a greater number of females in their territory and instead had more extra-territorial copulations. Whether these extra-territorial copulations are due to female choice or success in male competition is unknown.     

The adaptive significance of non-contact mate guarding by males of the dragonfly,Nannophya pygmaea Rambur (Odonata: Libellulidae)

InNannophya pygmaea, ovipositing females were frequently disturbed by conspecific males. Disturbed females often copulated with one of these males or flew away from the pool. Females which flew away from the pool due to male disturbance often returned later the same day and mated with different males. A territorial male would guard his ovipositing mate by hovering above her, presumably trying to prevent her from moving out of his territory. A non-territorial male would also guard his mate in a similar way, both at a vacant water area which was not occupied by any territorial males, or within the territory of a resident male. In addition, both territorial and non-territorial males chased intruding males in an attempt to prevent their mates from being stolen. Territorial males defended their mates better than non-territorial males. Both males and females often mated more than once in the course of a single day. Some territorial males copulated with a new female while another mate oviposited in their territories. This observation supported the “multiple mating hypothesis” proposed by Alcock (1979) and Uéda (1979) but other evidence suggested that this is an inadequate explanation for the non-contact guarding ofN. pygmaea.     

The mating system of the Mediterranean monk seal in the Western Sahara

The mating system of the Mediterranean monk seal was studied combining the use of diverse technologies. Sexual dimorphism in size was limited. Sexual activity was only observed to occur in the water. The different segments of the population segregated spatially: females, pups, and juveniles aggregated inside two main caves, whose entrances were controlled by a small number (2–3) of territorial males that defended aquatic territories situated at the very mouth of the caves. Other territorial males defended aquatic territories located further away (5–30 km). The tenure of aquatic territories was nonseasonal and spanned several years. Relatedness among pups belonging to the same cohort was low or null, indicating a low level of polygyny, which is not surprising for an aquatically mating phocid with a protracted reproductive season. However, in addition, genetic relatedness showed a remarkable temporal periodicity. These results in combination point to the existence of a complex social structure in this species.