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.     

Should I Stay or Should I Go Now: Late Establishment and Low Site Fidelity as Alternative Territorial Behaviors

In species in which males defend territories for breeding, males may differ in territorial behavior; alternative behaviors among territorial males are not well understood. In our long‐term study of partially‐migratory song sparrows, we have observed that most territorial males establish territories before females begin nesting and remain site‐faithful both within and between breeding seasons; however, some males establish territories later in the season (late establishers) and/or change territory locations either within or between seasons (movers). Whether late establishment or moving are equally successful strategies for territory defense, or best‐of‐bad‐job options, is not known. Here, we compare the frequencies of these behaviors to demographic variables over a 9‐yr period and compare lifetime tenure and early season nesting success for males who differ in site fidelity and timing of territory establishment. Across years, late establishing was negatively correlated with the return rate of previously territorial males; moving was positively correlated with the number of occupied territories at the start of the breeding season (territory density). While moving was independent of number of years on territory, late territory establishment only occurred in a male’s first year as a territory holder. Of 88 males, 25% established their first territory late, primarily in undefended space; 31% moved. Late and early establishers did not differ in lifetime tenure; movers, however, had longer lifetime tenure than site‐faithful males. Among early establishers, movers and non‐movers did not differ in the number of successful early nests/year or number of young fledged/year; among late establishers, however, movers had significantly higher early nesting success by both measures. Late establishers who moved had higher early season nesting success and higher early season nesting success/year than site‐faithful early establishers. Thus, individual variation in the timing of territory establishment and site fidelity may be facultative alternative territorial strategies.     

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.     

Accepting unrelated broods helps replacement male yellow-headed blackbirds attract mates

Replacement male yellow-headed blackbirds (Xanthocephalus xanthocephalus)did not destroy broods sired by the previous territorial maleand they showed no aggression toward females with unrelatedbroods. To test whether their tolerance of unrelated young wasmisdirected normal parental care, we removed males from experimentalterritories after primary nests were completed but before secondarynests were initiated. Replacement males fed young that theypresumably had sired in secondary nests and ignored foster youngin primary nests, whereas control males fed young in their primarynests. To identify potential benefits of accepting unrelatedyoung, we analyzed patterns of within-season breeding dispersaland of female settlement on territories following nest lossesto predators. Although some female yellowheads do renest onthe same territory following nest failures, the number of nestsinitiated on territories after a predation event was significantlylower than the number initiated on territories without predationover the same period of time. This implies that late-settlingfemales use the number of active or failed nests and/or thenumber of females on a territory when choosing where to breed.If replacement males that accepted unrelated offspring attractmore new females in the remainder of the current breeding seasonthan infanticidal males, then tolerance of unrelated young byreplacement males may be adaptive in some polygynous birds.     

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.     

Mating system and size advantage of male mating in the protogynous swamp eel Monopterus albus with paternal care

In fish with paternal care, protogynous sex change (female to male) is rare and has only been reported from species with haremic polygyny. The swamp eel, Monopterus albus, is a protogynous fish with paternal care, but little is known about its mating system. To understand protogyny in this species, we examined the mating system and male size advantage in mating in M. albus under semi-natural condition. Females swam over wide ranges and visited multiple male nests. Males defended a narrow territory around nests against other males that approached nests; at these nests, males courted and accepted visiting females. After spawning inside nests, caring males continued to perform courtship activities, and multiple breeding was observed. These observations suggest that the M. albus mating system is male-territory-visiting (MTV)-polygamy. Larger males had nests, and mated more frequently compared with small males. Because small initial males of this species are not found in nature, and because M. albus does not engage in sneaking tactics, larger nesting males do not suffer from reproductive parasitism. Thus, protogyny in this fish is likely consistent with the predictions of the size-advantage model. Biting attacks by territorial males of this predatory fish seriously wounded intruding males, occasionally resulting in the death of the intruder. We discuss the possibility that sexual differences in mortality rates in small fish may facilitate the evolution of protogyny in this species. Protogyny of the swamp eel is, to our knowledge, the first example of an MTV-polygamous mating system in a fish with paternal care.     

Resource defence on exploded leks: do male little bustards, T. tetrax, control resources for females?

In lekking species, female choice of particular males for mating is based on male characters that are not related to immediate gains such as access to resources. This is, however, unclear in exploded-lekking species, as male territories may contain resources for females; it is thus the degree to which males regulate female access to these resources to obtain matings that should distinguish lekking from nonlekking species. We investigated the relationship between resources for females, male distribution and male attractiveness in little bustards. One parameter determining resource quality for females was invertebrate availability: females laid eggs in grassland fields (grass, alfalfa and ryegrass) with significantly higher invertebrate biomass than others of similar cover but nests were in areas that did not have higher invertebrate biomass, at a scale of 1 km². Males set up territories and females laid eggs in areas that had a high proportion of alfalfa, but not high mean invertebrate biomass. Thus males set up territories at resources potentially used by females, but that appeared not to be critical for breeding. In addition, females did not use the amount of resources defended as a criterion for mate choice, as male attractiveness was not related to various territorial resources. Locating the lek in areas attractive to females might be a way of increasing the probability of encounter between the sexes. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

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.     

Habitat selection and polygyny in breeding Corn Buntings Miliaria calandra

Unlike many other polygynous passerine species, female Corn Buntings Miliaria calandra apparently do not suffer costs by pairing polygynously, yet it is unclear whether this is because polygynous males hold the highest quality territories or because pairing with polygynous males is unimportant in determining female reproductive success. Male Corn Buntings on North Uist, Scotland, consistently defended territories which contained nesting habitat, and females often foraged outside male territories when provisioning nestlings. Females showed strong preferences for nesting in uncultivated land, and 80% of nests were under Hogweed Heracleum sphondylium, possibly because this provided cover against predation and the weather. When provisioning nestlings, females showed strong preferences for foraging in cereal crops, probably because this habitat provided better food resources and/or better cover from predators. Males were unpaired or paired with one to three females per breeding season, but variation in territory size or vegetation composition did not explain differences in the number of females paired with individual males. We suggest that when females neither gain benefits nor suffer costs by breeding polygynously, and males do not differ greatly in the areas of habitat selected, polygyny can arise through random female settlement within the nesting habitat.     

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.     

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.     

Annual variation in the mating system of the dragonfly Paltothemis Kneatipes (Anisoptera: Libellulidae)

The libellulid dragonfly Paltothemis lineatipes exhibits annual variation in male mate-locating behaviour, variation that appears to be related to changes in the number of males competing in an area. In a year when males were numerous, as reflected in a high rate of male-male interactions, individuals defended small streamside territories, regularly raided neighbouring territories in an attempt to steal females there, and often were forced off their territory after a few hours of daily ownership. In this year, temporal partitioning of a given site was common, with the same males taking it in turns to defend the location for part of each day. In a year when males were much less numerous, as seen in a very reduced rate of aggressive encounters, males defended territories more than twice as large on average than in the high-density year; they rarely took females from neighbours, and they much less frequently 'shared' ownership of a site with another male on a given day. The changes between low- and high-density years were not associated with a change in average daily copulatory success of territorial males, although the variance in mating success was significantly greater in the high-density year.     

Female mate choice based on territory quality in barn swallows

Female mate choice based on territory quality is difficult to study because territories often contain many resources, which are difficult to quantify. Here, using the barn swallow (Hirundo rustica gutturalis) breeding at an outdoor breeding site in Japan, where each male defends only a small territory containing old nests, we studied whether females choose social mates based on territory quality. Since the territories of this species contain few other resources, territory quality can easily be assessed by quantifying old nests in the territory. We made the following four observations: (1) male swallows displayed old nests in their territories to females; (2) the old nests used for the first clutch were less broken than the other old nests within the same territory; (3) territory quality, defined by the number of old nests weighted by the intactness of each old nest, predicted the productivity of the territory; and (4) males with better territories paired with females earlier, and hence bred earlier, than those with inferior territories. The relationships remained significant even after controlling for male morphological traits. Based on these results, we can infer that female swallows choose their mates based, in part, on territory quality.     

Piracy mating by large males in a monogamous substrate-breeding cichlid in Lake Tanganyika

Telmatochromis temporalis is a biparental substrate breeding cichlid that mates in size-assortative monogamy at the southern end of Lake Tanganyika. In this population, at least a few months before the rainy season, large males (90–110 mm TL) did not pair. However, they maintained large territories that were defended against conspecific large males and that contained several pairing small males (50–80 mm). Large males occasionally visited the nests of the small pairs in their territories, and selectively took over the nests by attacking the owner male. During the takeover, the large males courted the small females in the nests and might have ejaculated sperm. After the large males left the nests, the small owner males conducted paternal care at the nests. These behaviour patterns of large males are similar to piracy mating reported in other polygamous fish species. The piracy mating of large maleT. temporalis seems to be an alternative tactic when large females are not available.     

Size-dependent use of territorial space by a rock-dwelling cichlid fish

Territoriality fundamentally influences animal mating systems and patterns of population structure. Although territory ownership is already known to contribute importantly to male reproductive success and the ecological coexistence of African rock-dwelling cichlids, the significance of variation in territory features has received little attention in these fishes. In Lake Malawi, males of Pseudotropheus tropheops "orange chest" defend territories on either of two substrate classes at Harbour Island: flat rock slabs lacking crevices and caves, or structurally complex boulder fields containing cave shelters. Focal watches of this species demonstrated that both territory size and occupancy on either substrate type depend on the size of male residents. Males larger than a threshold size exclusively held the largest and most structurally complex territories. After removal of conspecific residents, more vacant territorial areas on cave-containing substrate were reoccupied by "orange chest" males in full breeding coloration compared to vacant areas on flat substrate. These findings suggest competition among "orange chest" males for complex rocky substrate. Defense of caves was associated with enhanced male courtship rates: the number of caves within a male's territory was a better predictor of courtship activity than was male size or territory area. In addition to territories being crucial for male reproductive success and therefore likely playing a role in sexual selection, male-male competition for caves in rock-dwelling cichlids may be promoted by the ecological advantage of enemy-free space. Smaller "orange chest" males lacking caves tended to move into adjacent boulder fields in the presence of predators, particularly at night. In contrast, males defending caves were more likely to remain on their territories when nocturnal predators were present. The territorial behaviors of P. tropheops "orange chest" that we observed in situ provide an instructive natural framework for testing the roles of substrate and ecology in the mating systems of rock-dwelling cichlid fishes.     

Uncertainty in territory quality affects the benefits of usurpation in a Mediterranean wrasse

Individuals should defend sites when the expected benefits ofthe territory exceed the cost of defense. However, if territoryqulaity is unpredictable or difficult to assess, the expectedpattern of territorial behavior is less clear. In a Mediterraneanwrasse, Symphodus ocellatus, mating success is skewed with 2%of nesting males getting more than 20% of the spawning success.Yet, variation in mating success is not explained by any knownphysical characteristic of males or their territories. Instead,females prefer nests with a recent history of mating successbecause males are less likely to desert the offspring she leavesbehind. Thus, territory quality is transient and determinedby interactions between the sexes. I measured the frequencyof territorial takeovers and the uncertainty in mating successamong days at a nest. Observations indicated that S. ocellatusmales usurped their neighbor's successful nests when males wereunsuccessful and larger than their successful neighbor. Sitesthat achieved mating success had a significantly higher probability(0.84) of remaining sucessful between consecutive days thanunsuccessful territories had of becoming successful (0.30).Unsuccessful males obtained higher and more certain fitnessreturns if they usurped a successful neighbor's territory. Interactionswithin and between the sexes drive uncertainty in success, whichinfluences territorial behavior in this species.     

Feeding territoriality and the reproductive consequences in brown skuas<Emphasis Type="Italic"> Catharacta antarctica lonnbergi</Emphasis>

In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.Research licence: Umweltbundesamt Bonn 13.4-94003-1/5-7.     

5. GENERAL NOTES

Olive Thrushes Turdus olivaceus olivaceus in Grahamstown, South Africa, were present in their territories throughout the year. Mist-net captures showed no seasonal fluctuation in the population density of adults. The breeding season had a main peak from August to November and a minor peak in April. Song output was greatest at the beginning of the breeding periods, when territorial boundaries were most keenly contested. There was a positive correlation between seasonal variation in breeding intensity and song output. The adult survival rate was estimated at 80%, and 56% of the fledglings were alive in their natal territories at the age of independence. In this study area, only females built nests and brooded nestlings. Only males sang. Both males and females defended their territories.     

Flock, pair and group living lifestyles without cooperative breeding by Australian Magpies Gymnorhina tibicen

The social behaviour of Australian Magpies Gymnorhina tibicen was studied at Linton in the Manawatu district, New Zealand, from June 1978 to November 1982. Territorial magpies defended all-purpose areas averaging 5 ha in size, and there was no correlation between territory size and the number of occupants. The mean adult survival rate was 0.85, and average annual productivity was 0.96 juveniles per breeding female. Nomadic flocks formed in the spring and foraged on open, treeless pasture. They mainly comprised 1–3-year-old magpies which did not breed. Territories were defended by monogamous pairs, and by groups ranging from three to six individuals. Some territorial groups probably originated in the flock, and may therefore have consisted of unrelated birds. Other territorial groups formed when juveniles did not disperse away from the natal territory, and were called family groups. Non-breeders did not help at the nest in any group, and there was no difference in the production of young by females breeding in group-defended territories compared with those in pair-defended territories. The opportunity to disperse into non-territorial flocks may have liberated Australian Magpies from cooperation.     

Prey availability affects territory size,but not territorial display behavior,in green anole lizards

The availability of food resources can affect the size and shape of territories, as well as the behaviors used to defend territories, in a variety of animal taxa. However, individuals within a population may respond differently to variation in food availability if the benefits of territoriality vary among those individuals. For example, benefits to territoriality may differ for animals of differing sizes, because larger individuals may require greater territory size to acquire required resources, or territorial behavior may differ between the sexes if males and females defend different resources in their territories. In this study, we tested whether arthropod abundance and biomass were associated with natural variation in territory size and defense in insectivorous green anole lizards, Anolis carolinensis. Our results showed that both male and female lizards had smaller territories in a habitat with greater prey biomass than lizards in habitats with less available prey, but the rates of aggressive behaviors used to defend territories did not differ among these habitats. Further, we did not find a relationship between body size and territory size, and the sexes did not differ in their relationships between food availability and territory size or behavioral defense. Together, these results suggest that differences in food availability influenced male and female territorial strategies similarly, and that territory size may be more strongly associated with variation in food resources than social display behavior. Thus, anole investment in the behavioral defense of a territory may not vary with territory quality.