Family-based winter territoriality in western bluebirds, Sialia mexicana: the structure and dynamics of winter groups

Winter residency is characteristic of the majority of cooperatively breeding birds, but the composition and dynamics of winter groups have been examined in relatively few. In 1996-1998, we examined winter territoriality in the western bluebird, a year-round resident that shows a limited degree of helping behaviour in central coastal California, U.S.A. In spring, most western bluebirds breed as socially monogamous pairs, but a small proportion of pairs (3-16%) have additional breeding-age males helping at the nest, usually assisting parents or brothers. We found that year-round residents commonly wintered in family groups that defended territories similar to those used in spring. Winter groups had an even sex ratio and formed early in the autumn, when hatch-year birds dispersed. More females than males left their natal groups to be replaced by an influx of immigrant hatch-year birds. Winter groups typically consisted of breeders and one or two sons from the prior breeding season along with one or more immigrant females. A second period of dispersal occurred in spring when winter groups broke up and most birds other than the breeding pair left the winter territory. When they bred, yearling males and females often bred with unrelated individuals from their winter groups. Sons were more likely to remain on the study area as yearlings when they wintered with both parents than when they wintered with just one parent. We suggest that young males stay the winter due to benefits of remaining in family groups on mistletoe-based winter territories. Subsequent localized dispersal of sons then leads to opportunistic kin-based interactions later in life. Copyright 2001 The Association for the Study of Animal Behaviour.     

Male foraging avoidance in female feeding territories in a harem polygynous cichlid in Lake Tanganyika

We studied foraging site partitioning between the sexes in Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid with harem-polygyny. Females maintained small territories against heterospecific food competitors within large territories of males, foraging exclusively at the inner side of their own territories (foraging areas). Males fed as frequently as females in their own territories, but mostly outside female foraging areas, although they frequently entered female territories and repelled food competitors from the territories. Soon after removal of the resident females, however, harem males, as well as many food competitors, invaded the vacant territories and intensively devoured prey of female foraging areas. This indicates that although female foraging areas appear to contain more food than outside the areas, harem males refrained from foraging there when the resident females were present. We suggest that harem males will attempt to keep female foraging areas in good condition, whereby they may get females to reside in male territories and/or promote female gonadal maturation.     

The foraging ecology of feral goats

Feral goats (Capra hircus) were studied in the Mahoenui giant weta reserve, southern King Country, New Zealand, from March 1992 to February 1993. The reserve supports the main population of the undescribed Mahoenui giant weta (Deinacrida sp.). Gorse (Ulex europaeus) is the dominant woody browse plant in the reserve and provides protection, shelter and food for weta. The activities, foraging behaviour and diet of feral goats within the reserve were measured by direct observation and analysis of rumen contents. Measures of nutrient levels indicate that gorse is adequate for goat growth only during late spring/summer, and becomes a maintenance food at other times of the year. Feeding (grazing and browsing) was the dominant activity of adult feral goats in the reserve. Females spent more time feeding than males. Grazing and browsing changed seasonally for both sexes, with grazing generally decreasing from autumn to summer, and browsing increasing from summer to spring. In every season females spent more time grazing than males, but males browsed more than females. Greater use of browse by males may be an effect of the presence of females. Browsing of gorse by goats may not be an important influence on weta survival.     

Conflicting Territory Use in Males and Females of a Monogamous Ungulate,the Kirk's Dikdik

In monogamous mammals it is often unclear why males do not defend larger territories to attract more than one female. I investigated the territoriality of the monogamous Kirk's dikdik, Madoqua kirki, a dwarf antelope, in which food resources increase with territory size and some males defend enough resources for more than one female. Yet, all males are paired monogamously. When males were removed from small territories, their female partners spent more time outside of their territories than females in large ones. When females were removed, their male partners almost never left. Pairs in small territories spent more time together than pairs in large ones. Paired males left mostly together with their females, apparently not on their own initiative. Presumably because females in small territories left more often, their males spent more time outside in the female's company than males in large territories. I argue that males in smaller territories can keep better track of their females and that they can effectively reduce their females' time outside. Male intrusion pressure was unrelated to territory size, but it increased in the presence of unguarded females. If large territories decrease the ability to mate guard, and if unguarded females attract competing males, then defending large territories may be uneconomical, even it they could attract more than one female. On the other hand, territories must be large enough to satisfy the requirements of a single female.     

Sexual differences in food re-caching by New Zealand robins Petroica australis

We presented monogamous pairs of New Zealand robins Petroica australis with a superabundant supply of artificial prey and then observed birds re-cache previously hoarded prey. Males and females frequently relocated and redistributed previously hoarded prey. Both sexes also re-cached prey that was originally hoarded by their mate. However, re-caching rates of retrieved and pilfered prey differed between sexes. Female-made caches were re-cached at the same rate by both females and pilfering males, while male-made caches were re-cached more frequently by pilfering females. The re-caching of previously hoarded prey is consistent with the hypothesis that re-caching reduces theft. However, re-caching pilfered prey suggests that it might also play a role in cache theft strategies. Higher rates of re-caching by pilfering females cannot be explained by current hypotheses used to account avian re-caching behaviours. We suggest two new hypotheses, sexual dimorphism in spatial memory and indirect mate provisioning, which may help to explain the evolution of re-caching in New Zealand robins.     

Sex-specific aggression and sex ratio in wintering finch flocks: serins and siskins differ

Males are dominant over females in many bird species. This may lead to male monopolisation of resources whenever food is scarce or clumped and secondarily to lower female survival rates. As a result of the consequent male-biased sex ratio in the adult population, competition may arise either (1) between males and females, as males attempt to exclude females from feeding patches, or (2) between males because females do not pose a competitive threat. We recorded agonistic interactions between males and females in wintering foraging flocks of serins (Serinus serinus) and siskins (Carduelis spinus) to test for inter-specific differences. Most of the aggressive interactions in serins were between males and females, whereas in siskins they were between males. We also compared sex ratios for each species during the winter, determined from separate trapping efforts over an 11-year period, to test whether the direction of aggression by males (i.e. male/male; male/female) relates to variations in female survival rates. The proportion of females was smaller in winter than in autumn for serins, but differences in siskins were negligible. Results are interpreted in relation to the social organization displayed by both species studied.     

Spawning and biparental egg-care in a temperate filefish,Paramonacanthus japonicus (Monacanthidae)

Synopsis Reproductive habits of a temperate filefish, Paramonacanthus japonicus, were studied on a rocky reef at Tsuyazaki, Fukuoka, Japan, from 1989 through 1990. Males had territories of 30–70m² and defended them from conspecific males and potential egg predators such as another filefish, Stephanolepis cirrhifer. Egg masses were found on the sandy bottom in male territories. Individual discrimination of males and females occurring in three male territories revealed that males and females stayed in stable pairs during one month of observation in 1989. In these stable pairs, males fed only within their territories, but females occasionally foraged outside. The occurrence of egg masses within male territories and biparental egg care showed that fish were reproducing as monogamous pairs. Contrary to this, males tagged in 1990 changed their territories after the disappearance of females, and males and females mated polygamously. Spawning was observed only four times during the study period, between 1633 and 1754h. Prior to spawning, the female prepared a spawning bed on the sandy bottom. The male nuzzled the female and the pair spawned, touching their gonopores on the spawning bed. Spawning was very quick and took only 1–3 seconds. The adhesive eggs were spherical with a diameter of 0.56 mm. They were mixed with sand particles and formed a doughnut-shaped mass of about 4 cm in diameter. One egg mass contained 3300–3800 embryos of similar developmental stage, which hatched 2–3 days later. P. japonicus appears to be monogamous but may also practice polygamy when pair-bonds are unstable.     

Evolution of monogamy in seaside sparrows, Ammodramus maritimus: tests of hypotheses

Seaside sparrows (Ammodramus maritimus) are monogamous throughout their range. We used male-removal experiments, measurements of territory quality and nest placement in relation to territory boundaries, and observations on foraging behaviour and on intraspecific aggression to assess the applicability of pertinent hypotheses that could explain the occurrence of monogamy in this species. Since experimentally widowed females can rear about two-thirds as many young to independence as pairs, made help is advantageous but not necessary for female reproductive success. Female-female aggression is probably not important in maintaining monogamy, but we need to learn more about it. We induced experimental bigamy in seaside sparrows in New York, thus showing that males will accept second mates. Territory quality (suitable cover, nest-sites and food) is very heterogeneous in the sparrow populations. However, ‘polygyny thresholds’ evidently are not exceeded either because resources on salt marshes are not limiting (cover, nest-sites) or because females can nullify the effects of resource (food) disparity between territories by feeding at distant sites outside their mates' territories.     

The Effect of Male Incubation Feeding,Food and Temperature on the Incubation Behaviour of New Zealand Robins

Because of finite resources, organisms face conflict between their own self‐care and reproduction. This conflict is especially apparent in avian species with female‐only incubation, where females face a trade‐off between time allocated to their own self‐maintenance and the thermal requirements of developing embryos. We recorded incubation behaviour of the New Zealand robin (Petroica longipes), a species with female‐only incubation, male incubation feeding and high nest predation rates. We examined how male incubation feeding, ambient temperature and food availability (invertebrate biomass) affected the different components of females’ incubation behaviour and whether incubation behaviour explained variation in nest survival. Our results suggest that male incubation feeding rates of 2.8 per hour affect the female’s incubation rhythm by reducing both on‐ and off‐bout duration, resulting in no effect on female nest attentiveness, thus no support for the female‐nutritional hypothesis. The incubation behaviours that we measured did not explain nest survival, despite high nest predation rates. Increased ambient temperature caused an increase in off‐bout duration, whereas increased food availability increased on‐bout duration. While males play a vital role in influencing incubation behaviour, female robins attempt to resolve the trade‐off between their own foraging needs and the thermal requirements of their developing embryos via alternating their incubation rhythm in relation to both food and temperature.     

Plasticity in the mating system of the longnose filefish,Oxymonacanthus longirostris, in relation to mate availability

The mating system of the longnose filefish,Oxymonacanthus longirostris, was examined on coral reefs of Okinawa, Japan. This species has been shown previously to be monogamous. Fish were usually found swimming together in heterosexual pairs with the male and female sharing the same feeding territory. However, both monogamous and polygynous (bigamous) males were found in the present study. Polygynous males, which were larger than monogamous males, visited and stayed several minutes in turn with each female within the territories. Although most males were monogamous in the early breeding season, over 20% of males mated polygynously in the late breeding season. The adult sex ratio in the former was unbiased, but became slightly female-biased toward the end of the breeding season because of the higher disappearance rate of males. The higher disappearance rate may be due mainly to a higher mortality rate of males resulting from a greater deterioration of physical condition during the breeding season. Thus, the mating system varied with the change of the adult sex ratio. Plasticity in the mating system of this species may be the outcome of male mating tactic depending on local mate availability.     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

The longnose filefish,Oxymonacanthus longirostris, usually lives in heterosexual pairs, the male and female swimming together and sharing the same territory. Pair territoriality in the species was examined in detail in relation to sexual differences in territorial defense activities. Rigorous pair territoriality was maintained only during the breeding season, although pairs used their home ranges exclusively to a certain extent, during the non-breeding season. The frequency of aggression against other conspecific pairs in the breeding season was higher than in the non-breeding season. Agonistic interactions appear to be over both mates and food resources, the strict pair territoriality in the breeding season possibly being due to mutual mate guarding. In intraspecific aggressive interactions, males usually led their partner females when attacking intruders. The feeding frequency of males was much lower than that of females in the breeding season. Mate removal experiments indicated that females could not defend their original territories solitarily and their feeding frequency decreased. Conversely, males could defend territories solitarily without a decrease in feeding frequency. These results suggest that males contribute most to the defense of the pair territory, with females benefiting from territorial pair-swimming with their partner males.     

Seasonal variation in haematological parameters in male and female Tinca tinca

This study was designed to investigate any seasonal (spring, summer, autumn and winter) changes in haematological parameters in the blood of Tinca tinca measuring the number of red blood cells (RBC), haematocrit, white blood cells, and total plasma proteins.The results show significant changes in RBC and haematocrit in males comparing spring and summer with autumn and winter, whereas in females the RBC remained constant for all 4 seasons but the haematocrit decreased in autumn and winter compared to spring and summer. The white blood cells of male and female animals were significantly lower in spring and winter compared to summer and autumn. In male fish total protein contents significantly decreased in autumn and winter compared to spring and summer, whereas in females protein output significantly decreased in winter compared to the other seasons. The results indicate marked seasonal variation in the blood of male and female Tinca tinca. This variation may play a important protective role for the survival of the animals.     

Reproductive strategies of Diaeretiella rapae (Hymenoptera: Aphidiinae) during fluctuating temperatures of spring season in New Zealand

Reproductive activities of naturally occurring population of Diaeretiella rapae (M'Intosh) (Hymenoptera: Aphidiinae) on Myzus persicae were studied during spring season in New Zealand. The cabbage seedlings were highly infested with the aphids (272±25 individuals/plant) with about 30% on average parasitised by D. rapae. Rate of parasitism was positively correlated with the aphid density (R ²=0.64). Adult emergence from aphid mummies was 90%, with a higher frequency of females than males. Increase in female/male sex ratio was found between early and late spring (1.1 vs. 1.8). Mating in D. rapae occurred throughout the day. Nearly half of the mating pairs collected were male–male pairs, which were especially prevalent during morning and evening while more male–female mating pairs were found during midday. Male–male mounting was probably because of low number of virgin females in morning and evening populations. About 80% females were mounted by smaller size males. Male–male mounting is discussed in correlation with operational sex ratio of D. rapae population.     

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.     

Population ecology of Mus musculus on Mana Island, New Zealand

Feral house mice ( Mus musculus ) living on 217 ha Mana Island, New Zealand, with no mammalian predators, were snap-trapped and autopsied. A 7-month breeding season took the population from a spring low to extremely high density in autumn. Litters were largest in the middle of the breeding season, and significantly larger on Mana than on the New Zealand mainland. Litter size in early pregnancy was similar for young and old mice but more embryos were resorbed by old females. The breeding season ended in April when adult females stopped ovulating and young failed to mature. When the population declined over winter no animals bred, they all lost weight, and even previously mature males lost their reproductive ability. Mice continue to grow throughout life and become larger than mice in most populations on the New Zealand mainland. The regular and pronounced seasonal pulse in Mana's mouse population contrasts with longer-term fluctuations generally seen in mainland populations at lower density in indigenous forest. These differences may be explained by absence of predators, habitat features or lack of any chance to disperse on the island.     

The song structure and seasonal patterns of vocal behavior of male and female bellbirds (Anthornis melanura)

The bellbird (Anthornis melanura) is a honeyeater endemic to New Zealand, which uses song to defend breeding territories and/or food resources year round. Both sexes sing and the song structure and singing behavior have not yet been quantified. The number of song types, spectral structure, repertoire size, and singing behavior of male and female bellbirds was investigated for a large island population. Song types differed between the sexes with males singing a number of structurally distinct song types and females producing song types that overlapped in structure. Singing behavior also differed between the sexes; males often sung long series of songs while females sung each song at relatively long and variable intervals. Singing by both sexes occurred year round but the frequency of male and female singing bouts showed contrasting seasonal patterns. The frequency of female singing bouts increased as the breeding season progressed, whereas male singing bouts decreased. In contrast to almost all studied passerines, female bellbirds exhibited significant singing behavior and sung songs of complex structure and variety that parallel male song. These results provide a quantitative foundation for further research of song in bellbirds and in particular the function of female vocal behavior.     

Differences in the diet of the male and female tui (Prosthemadera novaeseelandiae: Meliphagidae)

Abstract

Tui are shown to be highly nectarivorous but observations show differences in degree of importance between the three food categories; nectar, fruit and arthropods, according to season. Foraging observations were significantly different for male and female tui during spring and especially during autumn, with males taking more nectar and females taking more arthropods.     

Monogamy in lizards

Monogamy is relatively rarely reported in taxa other than birds. The reproductive system of many lizard species appears to involve multiple mating partners for both the male and the female. However, short-term monogamous relationships have been reported in some lizard species, either where the male defends a territory that is only occupied by a single adult female, or where males stay with females for a period of time after mating, apparently to guard against rival males. There are a few reported cases of more prolonged monogamous relationships in lizards, with the Australian sleepy lizard, Tiliqua rugosa, the best studied example. Adult males and females of this species form monogamous pairs for an extended period before mating each spring, and they select the same partner in successive years. The paper reviews possible functions of monogamy in this and other lizard species, and suggests that the additional perspective from studying lizards may enrich our overall understanding of monogamous behaviour.     

Female choice and the benefits of mate guarding by male mallards

Pair formation and breeding in many species of waterfowl are separated both temporally and spatially. Most studies of female choice in this group have focused on male characteristics at the time of pairing, with less attention given to how mate choice affects breeding season outcomes. In this study I compared pairing success, male plasma testosterone level and mate-guarding ability of male mallards, Anas platyrhynchos, in two experiments. In the first experiment females and males were group housed with equal sex ratios, thus allowing all of these males to pair. At the same time, an equal number of males was housed in groups without access to females and remained unpaired. In this experiment testosterone levels of paired and unpaired males during autumn (baseline) and spring (breeding) did not differ, indicating that the process of pair formation and breeding does not cause elevated spring testosterone levels in males. However, testosterone did temporarily decrease in paired males during the winter (pair formation) season. In the second experiment groups were male biased, allowing only half of the males to pair. Here paired males had significantly higher testosterone levels than unpaired males during the breeding season, but not during the preceding autumn. Together the results of these experiments indicate that successful pair formation predicts but does not alter male testosterone level during the breeding season. I also found that females paired to males with high levels of testosterone were missing fewer feathers due to forced copulation attempts by nonmates, suggesting that females may choose males based on their mate-guarding abilities. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Mate-sharing costs in polygynous Northern Lapwings Vanellus vanellus

In this study bigamous female Northern Lapwings Vanellus vanellus received significantly less incubation relief from their males than monogamous females. On average, monogamous males spent 34.3% of their time incubating and bigamous males 29.9%. Bigamous males divided their effort between their nests, incubating on average 9.4% on primary nests and 20.5% on secondary nests. Bigamous females compensated for the lack of male relief. Primary females incubated for 71.8% of their time, secondary females for 64.2%, while monogamous females spent 52.7% of their time incubating. As a result, there was no significant difference in total nest attentiveness among nests of different status. Primary and secondary females received equivalent incubation relief from the male. Bigamous males increased their contribution to incubation significantly as the season progressed. A bigamous male's distribution of incubation relief between his females was unrelated to female body mass, or to the degree of asynchrony between primary and secondary females in arrival and laying. Incubation time was significantly, negatively, correlated with total nest attentiveness. Monogamous females spent most time, secondary females spent an intermediate time, and primary females spent the least time on maintenance behaviour (foraging, comfort behaviour, inactivity). No significant differences were found in hatching success among females of different mating status. However, the ratio of unhatched to hatched eggs (i.e. the eggs that remained in the nest at the time of hatching) differed significantly: secondary females hatched a smaller proportion of their eggs than monogamous and primary females.