Space use in a multi-male group of the group-living lizard

The aim of the study was to determine the space use of individuals in a multi-male group of the armadillo lizard Cordylus cataphractus . Specific focus was on the adult males and females of the group, because, ultimately, the manner in which adult males and females utilize space determines the mating system. A two-dimensional (2D) grid pattern was delineated at the crevice of a free-living, multi-male group to facilitate the recording of the positions of individuals during observations. All marked individuals ( n =55) that were visible were scanned through a telescope, and their respective positions were recorded at 30-min intervals over several days during the breeding season. Arcview 3.2 Geographical Information Software was used to create a computerized replicate of the 2D grid pattern at the crevice and to depict graphically the recorded positions for each individual using a minimum convex polygon procedure. From the polygons generated by this procedure, it was clear that the space use of adult males overlapped greatly with the space use of adult females, but that there was no overlap in space-use among adult males. Space use overlapped greatly among subadults and juveniles and also among these size classes and adult males and females. Adult male C. cataphractus are territorial, defending specific sites at the crevice. The territory of each male in the group incorporated at least one female. The space use of several females overlapped with the space use of two or more males. Males were observed to mate with one or more females, and one female was observed to mate with two different males on more than one occasion. Cordylus cataphractus males display typical territorial polygyny whereas females may be promiscuous.     

One Norwegian territory of the Marsh Tit Parus palustris during 35 years

One Norwegian territory was occupied by successive pairs of Marsh Tits Parus palustris during 35 breeding seasons. Only ten males and seven females were involved. Each pair showed strict site fidelity and kept to the territory all year round. The pair bond lasted throughout life, and the duration varied from one to eight successive years. The number of successive mates varied for the males from one to two females and for the females from one to four males. Assuming that the birds were juveniles or 1 year old when ringed, they survived, on the average, 4.4 years. The oldest male survived for at least 10 years 4 months and the oldest female for 8 years 4 months. The mean survival rate of adults from one breeding season to the next was 75.8%, which is remarkably high when compared with the genera! findings in tits.     

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.     

Intra- and Inter-seasonal Variation in the Socio-Spatial Behavior of Adult Male Collared Lizards, Crotaphytus collaris (Reptilia, Crotaphytidae)

When individuals maintain strong inter‐seasonal philopatry to the same territories, males may be able to re‐establish territory occupancy without intense intra‐sexual aggression, and instead spend more time courting females early in the reproductive season. Furthermore, when some males have prior experience defending the same territories, it may be necessary for young males to exhibit higher levels of aggression because they are establishing a territory for the first time. We tested these hypotheses by examining within‐season (1992 and 1997) temporal variation in the social behavior of adult male collared lizards of known age and prior territorial experience in a population where inter‐season philopatry to territories is high. Contrary to expectations, the frequency of aggression exhibited by males with and without prior territorial experience did not differ. The frequency of intra‐sexual aggression was higher in 1992 than in 1997, perhaps because male competitors were more abundant in 1992. Although there was an interactive effect of year, male display and patrol were low at the beginning of the reproductive season in Apr. and May, reached peaks during midseason in June, and then decreased as reproduction ended in July. The size of territories showed a similar pattern, with males defending larger areas in June. Our data support the philopatry hypothesis in that the establishment of territories occurred without high levels of aggression early in the season, perhaps because territory boundaries have been well defined by high rates of patrol and advertisement during the middle of the previous season. Inter‐sexual interactions were most frequent in June rather than at the beginning of the reproductive season. Adult females are producing their second clutches and yearling females are producing their first clutches in June. The high rate of inter‐sexual encounters in June supports the hypothesis that males allocate more time to courtship when females are receptive because there are more reproductively active females at this time. The temporal pattern of activities in adult Crotaphytus collaris appears to function as a compromise between competing intra‐ and inter‐sexual social demands on males, allowing males to maximize mating opportunities as well as maintain future access to productive territories.     

Female Choice Predicts the Best Father in a Biparental Fish,the Midas Cichlid (Cichlasoma citrinellum)

The Midas cichlid is a monogamous, biparental species. It breeds in a highly competitive system where pairs have a low probability of raising fry to independence. Both parents must cooperate to retain the territory and protect the fry from predation. Previous experiments showed that females prefer large, aggressive and sexually experienced males as mates but males do not display any consistent preferences. Here I present the results of two experiments designed to see whether qualities preferred by females correlate with increased success in retaining territories and in providing parental care. Pairs with either large or aggressive males had an advantage in appropriating and holding a breeding territory; reproductive experience conferred no advantage in usurping a territory. Aggressive and reproductively experienced males had an advantage in defending the brood from predators of fry, but size had no effect. Thus, the qualities preferred by females confer advantages both in holding territories and in protecting fry. In contrast, males need not be selective because females, once in possession of a brood, defend it equally well regardless of size, aggressiveness, or reproductive experience. The system is one of mutually enforced monogamy based on female choice; females drive the system because they provide more investment than do males (combining gametes and time) and because this investment is a reliable resource to the male.     

Spawning behavior of the collared knifefish, Cymolutes torquatus (Labridae) in Papua New Guinea

A colony of about 10 males and nearly 50 females of Cymolutes torquatus was found living over a sandy substrate on the northwest coast of Normanby Island, Papua New Guinea at depths 14.6 to 18.3 m. Each male appeared to maintain a harem territory, approximately 10 to 14 m², with 4 or 5 females living in sub-units of this territory. Eleven matings of 4 males with 2 to 4 females each were observed on 9 and 10 April 1993. The broadcast spawning of C. torquatus is similar to that of razorfishes in the genus Xyrichtys, but in contrast to most tropical sand-dwelling fishes which mate at dawn or dusk, C. torquatus mated mid-morning between 9:38 to 10:50 h. Sexual dichromatism, color changes during mating, and cleaning of C. torquatus by juvenile Halichoeres zeylonicus are documented with color photographs.     

Group living, breeding behaviour and territoriality in the Subdesert Mesite Monias benschi

In 1997–2000 we studied a population of Subdesert Mesites Monias benschi consisting of 35–68 adults comprising 32 groups of two to nine birds (modal group size of four). The study population was significantly male-biased in 1999 but not in 1997 or 1998. Overall, both sexes were philopatric, but when dispersal (or eviction) occurred, it appeared to be female-biased. Over 40% of groups contained more than two adult males, whilst < 15% contained more than two adult females. Whilst there was no evidence of behavioural dominance by females, intrasexual aggression within groups was observed only amongst females. In contrast to other birds occupying the same habitat, breeding in mesites was not tied to rainfall, and occurred throughout the year. Each breeding unit constructed several nests every year, only one of which was used. All adult males and at least one adult female co-operated to raise one or two clutches of one or two eggs per year. Males and females contributed equally to incubation. Chick production and chick survival were not related to group size or territory size. Groups defended large, permanent, and multipurpose territories and all group members contributed to territory defence. Territory size was positively correlated with the number of males in groups, but not with overall group size. Territories were tightly packed with very few areas unoccupied. Transect surveys conducted throughout the narrow geographical range of this species revealed its presence in a range of semi-arid habitat types. Small groups were more likely to be detected in intact, high-stature forest, whilst large groups were more likely to be detected in low-stature forest containing numerous spiny, xerophytic trees Didierea madagascariensis .     

Social organization in the pheasant (Phasianus colchicus): harem formation, mate selection and the role of mate guarding

Harem formation and mate selection were studied in the pheasant in order to determine the advantages of territorial harem defence polygyny to the two sexes. We investigated the factors affecting harem size and the advantage to a female in remaining with one territorial male during breeding.
Female group size declined during late March and early April as females moved from large overlapping ranges into smaller, more widely dispersed breeding ranges. The proportion of female groups accompanied by males increased during this period.
Some males had a disproportionate share of females. Settled females were monogamous but, because a female's nest was generally outside the male's territory, her home range was larger than his territory.
Harem members were usually from the same winter group. Harem size was not related to territory quality in terms of food supply or nesting cover. Females were loyal to one male in more than one year even if his territory position changed. Older, territory-owning males had more females, both adult and immature, than males with newly-established territories. Harem size was not correlated with territory size.
We conclude that the mating system of the pheasant is based on mate guarding which protects females not only from the risk of predation or injury, but also from excessive energy expenditure incurred through being chased by other males. When escorted by a territorial male, females spent three times as much time feeding, one-fifth as much time running, and one-tenth as much time alert, as they did when not guarded.     

The use of auxiliary courts by the lek-forming White-bearded Manakin Manacus manacus (Aves, Pipridae)

The White-bearded Manakin is a passerine specially noted for elaborate courtship. Each resident male has a cleaned oval court delimited by saplings in leks. No study mentions the use of more than one court by a territorial male during the breeding season. We report the use of auxiliary courts by males in the lowland forest of southeastern Brazil and discuss its probable function in attracting females for mating. Additionally, we experimentally modified a male’s main courts, testing that auxiliary courts serve as alternative display places. Twelve males from four different leks were observed for 145 h. Six males used from one to four auxiliary courts located 1.0 to 8.1 m from their main courts. The males that also used auxiliary courts displayed more than males that used only the main court. Nevertheless, the proportion of female visits per display time indicates that males that used only the main court have greater efficiency in attracting potential mates. Individual males responded differently to the experimental modification of their main courts, but one male avoided the modified court. The use of auxiliary courts may be a strategy adopted mainly by peripheral males to attract females that visit more successful males with central territories on the lek. In the short term, auxiliary courts function as optional display places in cases of loss of the main court. In the long term, the use of auxiliary courts may be involved in the temporal persistence of lek areas.     

The social and reproductive behavior of the eyed flounder,Bothus ocellatus,with notes on the spawning ofBothus lunatus andBothus ellipticus

Synopsis Observations of the Caribbean eyed flounder,Bothus ocellatus, were made during two field trips to Bonaire, Netherlands Antilles (January 1990, and December 1990 – January 1991) at depths from 3 to 6 meters. Data were collected by six scuba divers during 54 dives totaling 70 hours. Three mating groups were observed, each consisted of a male and 1 to 6 females. Each female had a distinct subunit within the male's territory. Courtship activity began about one hour before sunset and continued until sunset, when the male retired for the night. Both males and females retired into the sand at locations outside their daytime territory. Every day, males attempted to mate with each female individually within their territory. Spawning began with the male moving under the female who was resting on the sandy bottom. The pair then made a slow upward rise approximately 15 to 75 cm above the substrate, which culminated in a release of a cloud of gametes.Bothus lunatus andBothus ellipticus exhibited similar social organization toB. ocellatus, but differences in the spawning rise were observed. Intraspecific agonistic behavior was observed within males ofB. ocellatus andB. ellipticus.Senior author     

Turnover and dispersal in a Peregrine Falco peregrinus population

In south Scotland, most Peregrines returned to the same territories to breed in successive years, though a few females changed territory from one year to the next.
Annual mortality among breeding birds was at most 9% among females (or 11% in both sexes combined). There may have been considerable annual variation, however, and excluding one exceptional year out of five reduced the estimate for females to 7%. These estimates are maxima, but are still considerably lower than those obtained from ring recoveries of dead birds reported by members of the public.
Among trapped birds, four males first bred at age two years, one at three and another at four or five; two females first bred at one year, 13 at two years old and one at three. Five other females which were seen to be in first-year plumage but were not trapped, also laid eggs, and 12 other such paired females held territory but did not lay. Only one paired male held territory in first-year plumage.
In their movements between natal and breeding territories, some females moved further than males, with median distances of 83 and 58 km respectively. In addition, of birds trapped breeding in the study area, a greater proportion of the males than of the females had been born locally, despite an equal sex ratio among fledglings; this was also consistent with a greater dispersal of females. In general, Peregrines made much longer movements in their first year of life than subsequently. Movements were in any direction.     

Spacing pattern and body size composition of the protandrous anemonefish Amphiprion frenatus inhabiting colonial host anemones

The protandrous anemonefish Amphiprion frenatus often forms a group consisting of a large female, a small male, and a smaller nonbreeder at an isolated single host anemone, where home ranges of subordinates were covered with the female's home range. Within the group, the dominant individuals suppress the growth of subordinates, resulting in large size differences. The spacing pattern and body size composition of A. frenatus on colonial hosts were investigated in Ishigaki Island, Okinawa, Japan. Six breeding pairs and 14 nonbreeders inhabited a colony of 157 anemones. Each pair maintained a territory in which pair members used different hosts. Nonbreeders had unstable home ranges on the outskirts of or in the pairs' territories. Body size differences between males and females in pairs and between males and nonbreeders were small. The small size differences in the colony of hosts are caused by reduced suppression of growth of subordinates by the dominant individuals. The total area of host anemones largely affects spacing pattern and social suppression of the anemonefish.     

Patterns of territory settlement and consequences for breeding success in the Northern Wheatear Oenanthe oenanthe

We examined the pattern of territory settlement and its consequences for breeding success in the Northern Wheatear Oenanthe oenanthe on Bardsey Island, Wales, during the breeding seasons of 1991-93. Males returned earlier than females, and older males returned earlier than first-year breeders. Although their boundaries shifted between years, the general location of territories was consistent during the three-year study. There was a high degree of fidelity to area and territory between years for both sexes. The order of territory settlement, from which a territory rank was calculated, was highly consistent for males between years irrespective of individual settlement patterns and territory fidelity. Patterns of territory settlement were less predictable for females, although there was a significant correlation between the mean territory ranks of paired males and females. There was a male-biased sex ratio in each year, and between 5% and 26% of males remained unpaired throughout the breeding season. Male mating status and breeding success were dependent on arrival date, territory rank and breeding density. Early-arriving, usually older, males were able to settle on territories first and were more likely to pair, while later-arriving individuals were more likely to remain unmated. These effects were consistent between years, and consequently territories could be classified as either preferred (accounting for proportionately more breeding attempts) or non-preferred. Territory quality as opposed to individual quality appeared to explain much of the variation in breeding success, and both sexes benefited by breeding on preferred territories through enhanced breeding success and an increased probability that their offspring would be recruited to the population.     

Seasonal Changes of Vocal Rates and Their Relation to Territorial Status in Male Galápagos Sea Lions (Zalophus wollebaeki)

Male vocal displays play an important role in sexual selection through both male–male competition and female choice. While this is supported by numerous studies in birds, much less attention has been paid to the role of such displays in mammals. We investigated the function of vocal displays in a polygynous mammal, the Galápagos sea lion (Zalophus wollebaeki). In our study population a large proportion of mature males are unable to establish a territory, providing the opportunity to compare the vocal behaviour between territorial and non‐territorial males. We examined how seasonal patterns of vocalizations differed between territorial and non‐territorial males and how the number of females present in a territory influenced behaviour of territorial males. We found that territorial males vocalized at higher rates than non‐territorial males, and territorial males vocalized more at the onset than towards the end of territory tenure. During the onset of territory‐tenure vocalizations of territorial males were directed more often towards other males than females. Furthermore, we found that vocalizations of territorial males were not only given in male–male interactions, but were also actively directed towards females. Territorial males vocalized at higher rates when more females were present in their territory. Our results suggest that vocalizations are important in male–male interactions, are relevant in territorial disputes and are used as a proxy for females to assess a male’s quality.     

Bigamy or monogamy with maternal egg care in the triggerfish,Sufflamen chrysopterus

Reproductive behavior and mating systems of the triggerfish,Sufflamen chrysopterus (Balistidae), were studied on the fringing reef of Sesoko Island, Okinawa. Both males and females maintained territories against consexual adults, feeding on benthic animals within their own territories. Each male territory overlapped one or two female territories, with mating occurring between the cohabitants. The monogamous males were smaller and foraged more frequently than the bigamous ones, suggesting that the former allocated more energy to growth rather than to improving reproductive success. Pair spawning occurred around sunrise, females only taking care of the demersal eggs until hatching, which occurred around sunset of the same day. On spawning days females foraged less frequently than usual, but as frequently as males. Females spawned at intervals of 5–7 days, usually shifting sites within their territories. Thus both feeding and spawning sites were available for females within their territories, providing males with the opportunity to monopolize females by defending their territories.     

Influence of Age and Prior Experience on Territorial Behavior and the Costs of Defense in Male Collared Lizards

In polygynous species where males maintain strong interseasonal philopatry to the same breeding territories, older individuals have prior experience defending their areas, whereas younger individuals are defending a territory for the first time. Theoretical and empirical studies predict that under such conditions the cumulative costs of defense may be lower for older experienced males as a consequence of familiarity with environmental and/or social conditions in their particular local habitats. We used quantitative data from focal observational studies and introduction experiments to test the hypothesis that older collared lizard males (Crotaphytus collaris) with prior territorial experience (3 yr+ males) acquire larger territories allowing them to court more, different females, with greater frequency without incurring higher defense costs. Consistent with this hypothesis, 3 yr+ males controlled significantly larger territories and courted more females, without having significantly higher rates of territory patrol, frequencies of advertisement display, or aggressive interactions with same‐sex competitors. Moreover, the intensity of responses to size‐ and age/experience‐matched tethered intruders by 3 yr+ males was lower than that by 2 yr males in their first season of territory defense. Our results support the hypothesis that age/prior occupancy of territories lowers defense costs allowing males to defend larger areas and increase opportunities to court females, perhaps increasing mating opportunities. By contrast, 2 yr males may need to respond more aggressively to intruders because their ownership of territories is tenuous as a consequence of shorter territory occupancy.     

Space use in the root vole: basic patterns and variability

Home range characteristics and spatial organization of the root vole Microtus oeconomus were assessed by radio-tracking The aim of this paper is to test the assumption that each microtine species employs a certain modal spatial system around which there is considerable flexibility In the studied population the basic modes of spatial organization were female territoriality and male use of overlapping home ranges Female territories were significantly smaller and less diversified in size than male ranges The size of male home ranges was positively correlated with body mass Each female territory was overlapped by several male ranges, larger male ranges usually overlapped more female territories The above spatial organization suggests that the prevailing mating system in the root vole is promiscuity, and larger males have an access to more receptive females than have small-sized males
Departures from these basic modes were found among young mature females from summer cohort, forming clusters even at low density, and occasionally among females in locally crowded habitat patches, where they held overlapping home ranges A single male usually monopolized such a cluster, excluding other males from the area used by the 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).     

Nest and mate choice in the red bishop (Euplectes orix) : female settlement rules

We investigated female settlement in a colony of red bishops(Euplectes orix), a territorial and highly polygynous weaverbirdwidely distributed over sub-Saharan Africa. An earlier studyshowed that male reproductive success is mainly determinedby the number of nests a male builds in his territory, whichappeared to be a good indicator of male quality. Because malesprovide no parental care or food resources within the territory,females sharing a territory do not compete for material resourcesand might therefore be expected to settle preferentially interritories of males that build many nests to gain the possiblegenetic benefit of high-quality offspring. An analysis of femalesettlement, however, revealed that females did not show a preferencefor territories of males with many nests and that the distribution of female breeding attempts with regard to the number of vacantnests within a territory could be explained best by randomfemale settlement in 3 out of 4 years. Females settled moreoften than expected by chance (in 3 out of 4 years) in territoriesalready containing occupied nests, indicating that residentfemales did not discourage settlement of additional females.However, sharing a territory with other females might imposecosts in terms of an increased predation risk because nestsin territories that contained other nests with young sufferedfrom higher predation than nests in territories that did notcontain other nests with young. Females therefore might tradethe possible benefits of settling in territories of males withmany nests against the costs of sharing a territory with otherfemales. This might result in the mating pattern found withrandom female settlement and male reproductive success beingdirectly proportional to the number of nests built. We discuss possible implications of this mating pattern for sexual selectionon males.     

Responses in spatial organization to manipulations of the food resource in polygynandrous alpine accentors

The alpine accentorPrunella collaris is a territorial and polygynandrous mating species. The breeding unit is a group consisting of about seven members who share a group territory which contains all the resources necessary for living. Each female holds a small territory around her own nest within the larger group territory. Supplemental food in the form of millet seed was provided for a total of 23 group territories to test the hypothesis that the abundance and distribution of food influences the size of a female's territory and her dispersion pattern which, in turn, determine the mating system. Both males and females had regular access to the feeders, but females and groups provided with feeders did not have smaller territories than those without feeders. No groups were divided into two by offering extra food. Provision of extra food resulted in an increase of group size but did not alter the sex ratio. All members, including new settlers, were sexually active and the sexual relationships among the members of a group were still multiple matings. Food addition did not prompt the fed females to choose nest sites nearer the feeder. Nest sites were restricted to particular rocky slopes and non-vegetated areas which were invariably free from snow each year due to the effects of strong winds and the topography. Females showed a nest-site fidelity for successive years. The reason why supplemental food had no effect on the territory size or dispersion of female alpine accentors is discussed.