THE EFFECT OF AGE ON THE BREEDING BIOLOGY OF THE KITTTWAKE RISSA TRIDACTYLA

1. From 1954 to 1956 inclusive, the biology of individual marked Kittiwakes was studied at North Shields, Northumberland.
2. It was concluded that older Kittiwakes reacted to the breeding stimulus earlier, more intensively and with greater success than younger breeding birds.
3. Birds with previous breeding experience returned to the colony before birds breeding for the first time and these before non-breeders.
4. Before breeding started, birds which had bred previously spent more of their time at the colony than those about to breed for the first time.
5. Birds breeding for at least the second time laid the first egg 7·5 days earlier than those breeding for the first time.
6. Breeding started one day later for every four days the return to the colony was postponed.
7. Older breeding birds showed greater nest-site tenacity, laid larger clutches and had greater breeding success than younger birds.
8. The chicks in broods of two (but not of one) increased in weight more rapidly where the parents had previous experience.
9. Breeding Kittiwakes showed strong colony tenacity, but 24% of the marked non-breeding birds were subsequently seen in other colonies.
10. Over half the birds retained the same mate as in the previous year.     

The effect of food on laying date and clutch-size in Tengmalm's Owl Aegolius funereus

The breeding of Tengmalm's Owl Aegolius funereus was studied at Umeå, Sweden, during the 1984–85. Mean clutch-size was one egg larger in 1984 than in 1985 despite the later laying in 1984. The difference in clutch-size was related to a better food supply in 1984. Daily weight increase of females during the prelaying period showed a high negative correlation with laying date in 1985, and a high positive correlation with clutch-size independently of laying date in 1984–85. This suggested that food eaten before and during laying had a great and direct influence on both laying date and clutch-size. Many females increased in weight during laying and most others decreased only moderately (relative to egg weight), suggesting that body reserves were not a main source for egg production.
Late breeding females were provided with extra food during the prelaying and laying periods in 1985. Fed females weighed more, bred eight days earlier and laid one more egg than controls. At the same laying dates in mid season, and after heavy snow-fall, clutch-size and female weight were larger in the fed birds than in controls, but this was not so near the end of the laying season. Although the earliest of the fed late breeders weighed more, and probably were less restricted by food availability just before or during laying, they did not lay more eggs than did early breeders. This result suggested some limitation on clutch-size that could not be overcome by the supplementary feeding. Weights of females during laying did not show any consistent relationship with clutch-size during successive laying date intervals, suggesting that clutch-size was not directly related to body condition.     

Adjusting parental investment to changing environmental conditions: the effect of food ration on parental behaviour of the convict cichlid, Cichlasoma nigrofasciatum

Animals that care for their offspring may vary the amount of care provided for a particular brood in relation to environmental conditions. Food availablity is one factor that may affect the costs and benefits associated with parental investment. The convict cichlid, Cichlasoma nigrofasciatum, is a small, substrate-spawning cichlid from Central America. Both male and female provide parental care for eggs and fry. Paris were kept at one of three ration levels, high, medium or low. Time spent in parental fanning by females was positively related to ration. Males spent less time fanning than females and their parental behaviour varied considerably between individuals. Males on the high ration spent slightly more time fanning than those on the lower rations. The number of eggs produced per spawning and the post-spawning weight of both males and females were significantly and positively related to ration. Foraging frequency of both males and females was inversely related to ration. There was no significant effect of ration on the frequency of mouthing eggs and young or on intra-pair aggression. Eggs of low-ration fish hatched earlier than those of medium- and high-ration fish but there was no significant difference in the number of days that the young survived. These results suggest that the allocation of time and effort between parental and maintenance activities differs in relation to food supply. Parents may provide more care for the large brood produced when food is plentiful but place more emphasis on their own survival when food is short and broods are small.     

Effects of supplemental feeding and female age on timing of breeding in the Alpine Accentor Prunella collaris

Supplemental food in the form of millet seed was provided to 1-year-old and older (2:2-year-old) female Alpine Accentors Prunella collaris, and the effect on timing of breeding was studied over four breeding seasons in central Japan. Only 1-year-old females responded to the supplemental food: additional food advanced their settlement date, the onset of the development of a cloacal protuberance and laying date. In contrast, no effect of the experimental feeding was found on the timing of breeding in older females and they invariably bred earlier than 1-year-old females. Having settled on the breeding grounds, females developed the cloacal protuberance and retained it for about 1 month until the last egg in the first clutch was laid. The development period of the protuberance corresponded to the copulation period. Because the copulation period was not shortened by extra food and no significant difference occurred in either of the two age classes, it is necessary for earlier breeding birds to accumulate the nutrients needed to form the protuberance as well as the eggs. Although older females relied less on supplemental food, they were significantly heavier and formed the cloacal protuberance earlier than 1-year-old females. One-year-old females depended more heavily on supplemental food than did older ones and the duration between settlement date and the onset of development of the protuberance of 1-year-olds was reduced by additional food. These results raised the possibility that older females fly from their wintering grounds carrying some food reserves which they can use for forming the protuberance, while 1-year-old females collect the food needed for forming it on the breeding grounds.     

Selection of nest sites by a hole-nesting duck, the Goldeneye Bucephala clangula

Nest site preferences were examined for a population of Goldeneye Ducks breeding in nest boxes in Värmland, central Sweden. Some nest boxes were occupied more often than others even if females returning to the same nest box were excluded from the analysis. Nest boxes located higher up trees were occupied more often than those close to the ground and some spatial 'cluster groups' of boxes were occupied more often than others. Otherwise nest site prefernces were not related to any measured physical attributes of the boxes. Prefernces for nest boxes seemed to be based mainly on a tendency for females to select those that had been occupied by other females in the preceding year, especially if they had bred successfully. As a result of this, the occupancy of nest boxes was not random over years but rather progressed in a series of runs; a period of consecutive years in which a box was occupied was followed by a period of years in which it was empty.
There were reproductive consequences for these prefernces in that females occupying preferred boxes were less likely to lose their clutch to a predator. These females also bred earlier in the year and produced larger clutches and broods than females breeding in other boxes.     

Effect of late winter food addition on numbers and movements of snowshoe hares

Summary Peak (1980) and early decline (1981) populations of snowshoe hares were supplied with extra food in late winter to test the hypothesis that snowshoe hare populations are limited by food supply. Food supplemented populations increased through immigration in both years but the response was more pronounced in the early decline population. Animals supplied with extra food lost less weight, had higher survival in some cases, and males began to breed earlier. Immigrants to the food addition area were of two types: those that established home ranges on or near the area and those that spent only a brief time there before returning to their initial range. The possibility that these latter individuals were prevented from remaining on the food grid by residents is discussed. Results indicate that food supply is one factor that can limit peak and declining populations of snowshoe hares but the relation of spacing behaviour to food supply and numbers must also be considered.     

Attendance patterns and behaviour in relation to experience and pair-bond formation in the Wandering Albatross Diomedea exulans at South Georgia

Recruitment of Wandering Albatrosses Diomedea exulans to the breeding population at South Georgia took between 2 and 8 years after they first returned to their natal colony. In successive seasons, from first return to pairing, the date of arrival became earlier and the number of days spent ashore and the time spent interacting with other birds increased. Pairing birds arrived earlier and spent more time ashore than birds of similar experience which did not pair in that season. In the season following pairing they returned at the same time as breeding birds, but most did not breed; when ashore they spent much of their time alone or with their partner at the nest site. They left in mid-season before other non-breeders and bred the following season. Some birds accomplished this process by spending much time (50–60 days) ashore in two or three seasons but most birds spent a similar total time ashore spread over more seasons. Until the season prior to breeding, the number of birds of the opposite sex with whom interactions occurred was proportional to the amount of time spent ashore. There was, therefore, considerable scope for inter-individual assessment of potential partners before interactions were confined essentially to a single partner in the season before the first breeding attempt.     

Sex, age, experience and condition as factors affecting arrival date in prospecting common terns, Sterna hirundo

Several studies have shown that seabird colonies consist to a large extent of young nonbreeders (prospectors). These individuals appear at the colony later in the season than established breeders. The reasons for this late arrival have remained unclear in most cases, mainly because of technical difficulties in collecting sufficient data from nonbreeding individuals. We used a novel transponder system to identify remotely the members of a common tern colony, including nonbreeders, during eight breeding seasons and we combined the system with automatic balances. Ninety-two per cent of prospectors returned for the first time when 2 years old and 88.9% of recruits to the breeding population had spent at least one previous season at the colony as prospectors. In both sexes, most individuals prospected for one season, but more males than females prospected for more than one season, although a higher proportion of females started breeding without a previous prospecting phase. Terns arrived earlier in the season the older they were and the more experience of the colony they had, but experience proved to be more important than age. Prospectors gained about 3 weeks with a previous prospecting season whereas an additional year of age allowed birds to arrive only about 6 days earlier. Prospectors returning later in the season arrived with lower body masses. Males on average arrived earlier at the colony than females. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Pair bond and breeding success in Blue Tits Parus caeruleus and Great Tits Parus major

Data from 939 nests of the Blue Tit Parus caeruleus and 1008 nests of the Great Tit P. major from nestboxes provided in superabundance in mixed forest study sites between 1976 and 2001 were analysed to examine the effects of mate retention on breeding success and the relationship between mate fidelity and site fidelity. Most birds retained their former partner (76% in Great Tits and 65% in Blue Tits). The probability of a pair divorcing was affected by male age in Great Tits, divorce being more likely in pairs with first‐year males. Great Tit pairs breeding together for a second season bred earlier, but had no higher breeding success than pairs breeding together for the first time. In Blue Tits laying date and start of incubation tended to be earlier in pairs breeding together for a second season, but hatching and fledging dates were not earlier than in other pairs. Great Tit pairs breeding together for two consecutive seasons bred earlier in the second season than in the first, but breeding success did not differ significantly between years. In both species, breeding performance did not differ between pairs that divorced after a season and pairs that stayed together. Thus breeding success did not determine whether a pair divorced or bred together again. Neither Blue Tits nor Great Tits improved their breeding performance through divorce. Blue Tit females even had fewer fledglings in the year after divorce than in the year before. Mate retention affected breeding site fidelity. Blue Tit females had greater breeding dispersal distances between consecutive years when re‐mating than when breeding again with the same mate. In Great Tits both males and females dispersed more when re‐mating than when retaining the former partner, suggesting that mate retention increased the chance of retaining the breeding site. In both species, breeding dispersal distances did not differ between pairs that divorced and pairs in which one mate disappeared. Because no major advantage of mate retention was evident, we suggest that mate retention evolved under different conditions than those found in study sites with high breeding densities and a superabundance of artificial nesting sites.     

The effect of supplemental food on the growth rates of neonatal,young, and adult cotton rats (Sigmodon hispidus) in northeastern Kansas,USA

In food-limited populations, the presence of extra food resources can influence the way individuals allocate energy to growth and reproduction. We experimentally increased food available to cotton rats (Sigmodon hispidus) near the northern limit of their range over a 2-year period and tested the hypothesis that seasonal growth rates would be enhanced by supplemental food during winter and spring when natural food levels are low. We also examined whether additional food resources were allocated to somatic growth or reproductive effort by pregnant and lactating females. The effect of supplemental food on growth varied with mass and season, but did not influence the growth rates of most cotton rats during spring and winter. In winter, small animals on supplemented grids had higher growth rates than small animals on control grids, but females in spring had lower growth rates under supplemented conditions. Growth rates of supplemented cotton rats were enhanced in summer. Northern cotton rat populations may use season-specific foraging strategies, maximizing energy intake during the reproductive season and minimizing time spent foraging in winter. Adult females invest extra resources in reproduction rather than in somatic growth. Pregnant females receiving supplemental food had higher growth rates than control females, and dependent pups (≤ 1 month of age) born to supplemented mothers had higher growth rates than those born to control mothers. Increased body size seems to confer an advantage during the reproductive season, but has no concomitant advantage to overwinter survival.     

Food supplementation and possible mechanisms underlying early breeding in the Florida Scrub-Jay (Aphelocoma coerulescens)

Food supplementation studies demonstrate the importance of resources in the timing of reproduction. Studies of Florida Scrub-Jays (Aphelocoma coerulescens) found that supplemented jays bred earlier than unsupplemented jays and that protein may play a critical role. In this study, free-living scrub-jays were provided with supplemental diets high in fat and protein (HFHP) or high in fat and low in protein (HFLP). Jays in both treatments bred earlier than unsupplemented controls (CNT), but HFHP-supplemented jays bred earlier than HFLP jays. To assess possible mechanisms, we measured testosterone (T) in males, estradiol (E2) in females, and corticosterone (CORT) in both. HFHP males had higher T than HFLP and CNT males, but treatment did not affect E2 levels of females. Pilot studies of scrub-jays in suburban environments suggest that the spatial and temporal predictability of food may influence corticosterone (CORT) levels. Suburban jays have year-round access to human-provided foods and breed earlier than wildland jays; thus, we compared CORT in all treatments in the natural site (wildlands) with those of suburban jays. CORT levels of suburban jays were lower than HFLP, HFHP, and CNT jays. HFHP-supplemented jays had lower CORT levels than those of HFLP and CNT jays. The observed differences in the timing of breeding, both between suburban and wildland populations and between experimental groups in the wildlands, may result from differences in the spatial and temporal predictability of food, and the nutritional differences in diets. Because CORT can negatively affect the reproductive axis, we postulate that nutrient availability, the predictability of food, CORT levels, and initiation of reproduction are inextricably linked.     

Relocation, reproduction and remaining alive in the orb-web spider

When mortality is high, animals run a risk if they wait to accumulate resources for improved reproduction so they may trade-off the time of reproduction with number and size of offspring. Animals may attempt to improve food acquisition by relocation, even in 'sit and wait' predators. We examine these factors in an isolated population of an orb-web spider Zygiella x-notata . The population was monitored for 200 days from first egg laying until all adults had died. Large females produced their first clutch earlier than did small females and there was a positive correlation between female size and the number and size of eggs produced. Many females, presumably without eggs, abandoned their web site and relocated their web position. This is presumed because female Zygiella typically guard their eggs. In total, c . 25% of females reproduced but those that relocated were less likely to do so, and if they did, they produced the clutch at a later date than those that remained. When the date of lay was controlled there was no effect of relocation on egg number but relocated females produced smaller eggs. The data are consistent with the idea that females in resource-poor sites are more likely to relocate. Relocation seems to be a gamble to find a more productive site but one that achieves only a late clutch of small eggs and few achieve that.     

Factors determining the laying date of the Waved Albatross Diomedea irrorata

The Waved Albatross laying period is between mid-April and the end of June. This paper investigates the factors determining when, during this period, a female lays. It is shown that birds breeding for the first time lay relatively late. For older birds, the laying date depends on date and outcome of the previous year's attempt: females that bred successfully in the previous year lay slightly later than in that year, while birds that failed lay earlier. A corresponding difference exists in the dates of arrival at the breeding grounds of successful and failed males. It is argued that one calendar year is not quite sufficient for pairs that have reared a chick successfully to regain breeding condition, while failed breeders can choose the laying date more freely.
Under catastrophic El Niño weather conditions in 1983, all albatrosses arrived late at the breeding grounds, but still the birds that had failed in 1982 were clearly earlier than those that had reared a chick.     

油茶地蜂生活史及相关生物学习性

油茶地蜂Andrena camellia Wu 是江西、湖南等地油茶花期主要传粉昆虫之一。2006–2007年在江西宜春对油茶地蜂除成虫外各虫态的形态特征、生活史及幼虫和成虫的行为等生物学特性进行了观察研究。结果表明： 该蜂在江西宜春1年发生1代。成虫10月中旬开始出巢活动, 雄蜂早于雌蜂3 d左右出现,完成交尾后数日死亡,雄蜂出巢后寿命18 d左右,雌蜂约38 d。卵期约8 d。幼虫期24 d左右,幼虫取食完花粉球后长成老熟幼虫进入滞育期,滞育期约241 d。翌年9月中旬开始化蛹,蛹期约30 d。出土后的雌蜂当天就可以交配,交配后的雌蜂在羽化地点附近选址筑巢,采集油茶的花粉和花蜜于虫室中制成球形蜂粮并在上面产卵,每粒花粉球上产卵一枚。     

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.     

捕食风险对不同饥饿状态下艾虎取食行为的影响

在室内条件下,将大鵟作为艾虎的天敌动物,通过双通道选择实验确定6 只成体艾虎在3 个捕食风险水平和4 种饥饿状态条件下的取食行为,探讨艾虎在取食过程中对饥饿风险与捕食风险的权衡策略。研究结果表明:在无捕食风险存在时,艾虎被剥夺食物0 d 和1 d 后对食物量不同的两个斑块中的取食量和利用频次均无明显不同(P > 0. 05),但对高食物量斑块的利用时间均明显高于低食物量斑块的(P <0.05),而艾虎被剥夺食物2 d和3 d后对高食物量斑块中的取食量和利用时间均明显高于低食物量斑块中的(P < 0.05),但在利用频次上均无明显差异(P > 0.05)。在面临低风险时,艾虎在4 种饥饿状态下均只利用无天敌动物存在的低食物量斑块,而基本不利用有天敌动物存在的高食物量斑块。在面临高风险时,艾虎不得不利用有天敌动物存在的食物斑块,被剥夺食物0 d 时艾虎对无风险、无食物量斑块的利用时间基本相同于对高风险、有食物量斑块的利用时间(P>0.05),而被剥夺食物1d、2 d 和3 d 后艾虎对高风险、有食物量斑块的利用时间明显高于无风险、无食物量斑块的(P< 0. 05)。在相同风险条件下,随着饥饿程度增加,艾虎在斑块中的取食量均明显增加(P< 0.05),而对斑块的利用时间和利用频次明显降低(P<0.05)。在相同的饥饿状态下,不同风险水平时,艾虎在斑块中的取食量无明显的差异(P>0.05),但在低风险和高风险时对斑块的利用时间和频次均明显低于无风险时的(P <0.05)。以上结果说明艾虎能够根据食物摄取率和自身的能量需求在捕食风险和饥饿风险之间做出权衡,当饥饿风险小于捕食风险时,艾虎趋于躲避捕食风险,当饥饿风险大于捕食风险时,艾虎趋于面对捕食风险,所采用的取食策略是减少活动时间和能量消耗,最大程度地提高单位时间内获得的能量。     

Reproductive biology of captive alpine ibex (Capra i. ibex)

We studied the reproductive biology of alpine ibex bred at the Peter & Paul Wildlife Park, St. Gallen, Switzerland, from 1930 through 1943 and 1968 through 1983. Peter & Paul Wildlife Park had supplied most of the original stock used to reestablish ibex populations in the European Alps after they had become extinct. Ibex females were seasonally polyestrous. Their estrous cycle averaged 20 days, and their gestation period averaged 167 days. Females between 1 and 15 years of age on average produced 0.78 young per year; females between 3 and 13 years of age produced 0.99 young per year. The average age at which captive ibex females gave birth was 8.44 years. Sex ratio at birth was unbiased. More singletons than twins were born, but the proportion of twins born during the years 1968 through 1983 was higher than during the earlier study period. There were no differences in the survival rates of male or female and single or twin offspring.     

Larval sensory abilities and mechanisms of habitat selection of a coral reef fish during settlement

In most animals, siblings from a given reproductive event emerge over a very short period of time. In contrast, many species of birds hatch their young asynchronously over a period of days or weeks, handicapping last-hatched chicks with an age and size disadvantage. Numerous studies have examined the adaptive significance of this atypical hatching pattern, but few have attempted to explain the considerable intrapopulation variation that exists in hatching asynchrony. I explored proximate determinants of hatching asynchrony by monitoring 112 Burrowing Owl (Athene cunicularia) nests in the grasslands of southern Saskatchewan, Canada, over 4 years. Age disparities between first- and last-hatched siblings (i.e., hatching spans) varied considerably, ranging between 1 and 7 days (mode = 4 days). These hatching spans increased with increased hatching success. Hatching spans also increased with larger clutches, but the increase was less than predicted given the increased time required to lay more eggs. Hatching span was unrelated to number of prey cached in the nest during egg laying (an index of food availability), and was unaltered by a year of super-abundant prey. Furthermore, pairs given extra food during laying had hatching spans equal to those of unsupplemented control pairs. These results were inconsistent with both the energy constraint and facultative manipulation hypotheses, which predict that hatching asynchrony should vary with the level of food during laying, when incubation onset is determined. Burrowing Owls were apparently free of food limitation early in breeding, yet may not have been able to optimize hatching spans because food conditions during laying were largely unrelated to food conditions during brooding. Thus, one of the premises for facultative manipulation of hatching asynchrony—that laying females are able to forecast post-hatch food conditions—may not have been met for this population of Burrowing Owls.     

Characteristic features of the reproduction of Koshima monkeys,Macaca fuscata fuscata: A summary of thirty-four years of observation

The reproductive data for Japanese monkeys,Macaca fuscata fuscata, which had been recorded for the 34 years from 1952 to 1986 on Koshima, were analyzed in terms of the influence of changes in artificial food supplies, the differences in reproductive success between females, the timing of births, and the secondary sex ratio. Koshima monkeys increased in number until 1971 when the population density was still small and artificial provisioning was copious. As described byMori (1979b), the severe reduction in artificial food supplies, which began in 1972, had an enormous deleterious effect on reproduction: the birth ratio of adult females of 5 years of age or more fell from 57% to 25%; the rate of infant mortality within 1 year of birth rose from 19% to 45%; primiparous age rose from 6 to 9 years old on average; and there was an increased death rate among adult and juvenile females. The prolonged influence of “starvation” may be seen in the significantly delayed first births of those females that were born just before the change in food supplies. When reproductive parameters are compared between the females who belonged to six lineages in the group during these periods, they were found to be rather consistent, although some individual differences can be recognized among females and subgroups. The apparent trend was that some of the most dominant females retained superior reproductive success while that of the second-ranked females has tended to diminish over the years since 1972. Such opposing trends were seen only in the most dominant lineage group and such a difference was not recognized among the females of other lineages. The difference in reproductive success is discussed in relation to both the different situations that arise because of the artificial food supplies and differences in feeding strategies. Multiparous females, after a sterile year, gave birth somewhat earlier than those who reared infants in the preceding year and, when artificial provisioning was intense, they tended to give birth a little earlier than during other periods. There is some evidence that the mortality of later-born infants was higher than that of earlier-born infants after 1972. However, this difference may not be responsible for the differential reproductive success of females since the timing of births did not differ among lineages. Furthermore, during the time when many females gave birth continuously, prior to 1972, the infant mortality did not differ with respect to the timing of births. The differences in infant mortality were not correlated with the reproductive history, parity or age of the mother, or with the sex of the infant. The secondary sex ratio varied by only a small amount, from slightly male-biased ratio (114: 100) when correlated with reproductive history, parity, age of mother, sex and survival ratio for preceding infants, timing of birth, and lineage of the female. Furthermore, the change in artificial food supplies did not cause any modifications of the secondary sex ratios, despite its enormous deleterious effect on reproduction. The secondary sex ratio of Japanese monkeys may not be influenced by the social factors mentioned.     

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.