FORAGING BEHAVIOUR OF NONBREEDING IMPERIAL CORMORANTS AT THE PRINCE EDWARD ISLANDS

Cooper, J. 1985. Foraging behaviour of nonbreeding Imperial Cormorants at the Prince Edward Islands. Ostrich 56: 96–100.

The Imperial Cormorant Phalacrocorax atriceps is an inshore, primarily solitary forager in the nonbreeding season at the Prince Edward Islands. All girds foraged within 400m of the coast, most seaward of a thick Macrocystis kelp bed. Mean dive duration was 40,2s with a maximum of 88s. The ratio of dives to surface rests was 2,68. Birds foraging very close inshore in shallow (<2 m) water dived for shorter periods than those farther offshore in deeper water. The Imperial Cormorant at the Prince Edward Islands appears to be rimarily a bottom-feeder. Foraging bouts were of short duration and little time was spent flying luring foraging. Imperial Cormorants dive for longer periods than do nearly all other species of cormorants.     

Does Foraging Performance Change with Age in Female Little Penguins (Eudyptula minor)?

Age-related changes in breeding performance are likely to be mediated through changes in parental foraging performance. We investigated the relationship of foraging performance with age in female little penguins at Phillip Island, Australia, during the guard phase of the 2005 breeding season. Foraging parameters were recorded with accelerometers for birds grouped into three age-classes: (1) young, (2) middle age and (3) old females. We found the diving behaviour of middle-aged birds differed from young and old birds. The dive duration of middle age females was shorter than that of young and old birds while their dive effort (measure for dive and post-dive duration relation) was lower than that of young ones, suggesting middle-aged birds were in better physical condition than other ones. There was no difference in prey pursuit frequency or duration between age classes, but in the hunting tactic. Females pursued more prey around and after reaching the maximum depth of dives the more experienced they were (old > middle age > young), an energy saving hunting tactic by probably taking advantage of up-thrust momentum. We suggest middle age penguins forage better than young or old ones because good physical condition and foraging experience could act simultaneously.     

Sex and individual differences in foraging behavior of Japanese cormorants in years of different prey availability

Japanese cormorants, Phalacrocorax capillatus, are sexually dimorphic seabirds with males that are heavier and that dive deeper than females. Sex differences in prey composition and foraging behavior of those rearing chicks at Teuri Island, Hokkaido, were examined by collecting food-loads from parents in 1992–1998 and by radio-tracking ten birds each in 1997 and 1998 when prey availability was different. Males fed more on benthic and epibenthic fishes (82% mass) than did females (34%) while females fed more on epipelagic and coastal fishes (53%) than did males (18%). Males made longer dives (53 s) at feeding sites closer to the island (7 km) than females (39 s, 13 km) in 1997. In 1998 when the availability of epipelagic fish seemed to be higher, there were no sex differences in dive duration and distance to the feeding sites (35 s and 9 km for males, 36 s and 10 km for females). This sex difference in foraging behavior with a poor availability of epipelagic fish suggests that high diving ability possibly enables males to feed on demersal fish. Birds specializing in coastal shallow waters around the island made long dives; hence they were probably foraging in bottom layers. Those foraging in deeper shelf waters made short dives and they were thought to forage in surface layers. Electronic Publication     

Sexual,Seasonal and Group Size Differences in the Allocation of Time between Vigilance and Feeding in the Greater Rhea,Rhea americana

We studied the effect of sex and group size on the proportion of time a greater rhea, Rhea americana, allocates to vigilance and feeding during the breeding and the non-breeding seasons. We analysed 175 records of focal animals that were feeding alone or in groups of 2 to 26 birds. In both seasons, males spent more time in vigilance and less time in feeding than females. Both sexes spent more time in vigilance and less time in feeding during the breeding season. Sexual and seasonal differences in vigilance were the result of different mechanisms. Males had shorter feeding bouts than females but there were no sexual differences in the length of the vigilance bouts. On the contrary, seasonal differences were the result of males and females having longer vigilance bouts during the breeding season but there were no seasonal differences in the length of the feeding bouts. During the non-breeding season, individual vigilance was higher in rheas foraging alone than in groups. In this case, solitary birds had longer vigilance and shorter feeding bouts than birds foraging in groups. We discuss the possible effect of intragroup competition and food availability on the allocation of time between feeding and vigilance in this species.     

Dive bout organization in the Chinstrap penguin at seal island, antarctica

Diving behavior of 2 breeding Chinstrap penguins (Pygoscelis antarctica) was studied focusing first and primarily on dive bouts rather than dives themselves. Analysis of dive bout organization revealed (1) though there are differences between solitary dives and dive bouts in dive duration and dive depth, the first dives of dive bouts do not differ from solitary dives in the dive parameters, (2) mean dive duration during bout correlates positively to both mean dive depth during bout and mean surface interval during bout, while number of dives during bout negatively correlates to both cost (consumed energy) and duration of a dive cycle during bout. These findings suggest the following possibilities on foraging behavior of penguins: (1) their decision to repeat diving depends on the result of the first dive at a site, and the first dives of bouts would tend to be searching or evaluating dives though they would be also successful foraging dives, (2) they repeat diving at a foraging patch until foraging efficiency decrease to a threshold of diminishing returns.     

Diving in shallow water: the foraging ecology of darters (Aves: Anhingidae)

Diving birds have to overcome buoyancy, especially when diving in shallow water. Darters and anhingas (Anhingidae) are specialist shallow-water divers, with adaptations for reducing their buoyancy. Compared to closely-related cormorants (Phalacrocoracidae), darters have fully wettable plumage, smaller air sacs and denser bones. A previous study of darter diving behaviour reported no relationship between dive duration and water depth, contrary to optimal dive models. In this study I provide more extensive observations of African darters Anhinga melanogaster rufa diving in water<5 m deep at two sites. Dive duration increases with water depth at both sites, but the relationship is weak. Dives were longer than dives by cormorants in water of similar depth (max 108 s in water 2.5 m deep), with dives of up to 68 s observed in water<0.5 m deep. Initial dives in a bout were shorter than expected, possibly because their plumage was not fully saturated. Dive efficiency (dive:rest ratio) was 5–6, greater than cormorants (2.7±0.4 for 18 species) and other families of diving birds (average 0.2–4.3). Post-dive recovery periods increased with dive duration, but only slowly, resulting in a strong increase in efficiency with dive duration. All dives are likely to fall within the theoretical anaerobic dive limit. Foraging bouts were short (17.8±4.3 min) compared to cormorants, with birds spending 80±5% of time underwater. Darters take advantage of their low buoyancy to forage efficiently in shallow water, and their slow, stealthy dives are qualitatively different from those of other diving birds. However, they are forced to limit the duration of foraging bouts by increased thermoregulatory costs associated with wettable plumage.     

Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua

We present data on diving pattern and performance (dive depth, duration, frequency and organization during the foraging trip) in gentoo penguins Pygoscelis papua , obtained using time-depth recorders ( n = 9 birds, 99 foraging trips). These data are used to estimate various parameters of foraging activity, e.g. foraging range, prey capture rates, and are compared in relation to breeding chronology. Foraging trip duration was 6 h and 10 h, and trip frequency 1.0/day and 0.96/day, during the brooding and creche periods, respectively. Birds spent on average 52%of each foraging trip diving. Dive depth and duration were highly bimodal: shallow dives (< 21 m) averaged 4 m and 0.23 min, and deep dives (> 30 m) 80 m and 2.5 min, respectively. Birds spent on average 71%and 25%of total diving time in deep and shallow dives, respectively. For deep dives, dive duration exceeded the subsequent surface interval, but shallow dives were followed by surface intervals 2–3 times dive duration. We suggest that most shallow dives are searching/exploratory dives and most deep dives are feeding dives. Deep dives showed clear diel patterns averaging 40 m at dawn and dusk and 80–90 m at midday. Estimated foraging ranges were 2.3 km and 4.1 km during the brood and creche period, respectively. Foraging trip duration increased by 4 h between the brood and creche periods but total time spent in deep dives (i.e. time spent feeding) was the same (3 h). Of 99 foraging trips, 56%consisted of only one dive bout and 44%of 2–4 bouts delimited by extended surface intervals > 10 min. We suggest that this pattern of diving activity reflects variation in spatial distribution of prey rather than the effect of physiological constraints on diving ability.     

Women from Venus,men from Mars: inter‐sex foraging differences in the imperial cormorant Phalacrocorax atriceps a colonial seabird

Colonial seabirds are central place foragers and likely to be subject to substantial competition for resources. Mechanisms proposed for reducing intra‐specific competition include differential inter‐sex area use mediated by adult choice. We used GPS loggers and dive recorders to study area use and dive depth in a total of 27 male and 26 female imperial cormorants Phalacrocorax atriceps breeding at a colony of some 6500 birds at Punta Leon, Chubut, Argentina during 2004 and 2005. Although time spent travelling and distances between the colony and foraging sites were similar for both sexes, males and females travelled away from their colony using routes virtually perpendicular to each other so that their foraging areas were distinctly different; females hunted close to the coast while males foraged offshore in deeper water. Consideration of foraging efficiency underwater, defined as the duration spent on the bottom divided by the dive cycle duration, showed that females were more efficient at depths < 40 m while males more efficient at depths > 40 m. We suggest that the substantial sexual dimorphism in this species may be responsible for the different depth‐linked foraging efficiencies and that selection for appropriate depths could lead to differential habitat use and putative differences in prey selection.     

BIOLOGY OF THE BANK CORMORANT,PART 1: DISTRIBUTION,POPULATION SIZE,MOVEMENTS AND CONSERVATION

Cooper, J. 1981. Biology of the Bank Cormorant, Part 1: Distribution, population size, movements and conservation. Ostrich 52: 208–215.

The Bank Cormorant Phalacrocorax neglectus is a marine species, endemic to southern Africa. Its non-breeding range extends from Walvis Bay to Cape Agulhas. Breeding range extends from Hollamsbird Island to Quoin Rock. Its distribution is broadly similar to that of kelp beds Ecklonia maxima. A total of 44 breeding localities supports approximately 18 000 adult birds; 12 800 (71%) occur on two islands (Ichaboe and Mercury) north of large kelp beds. Adult Bank Cormorants are resident but juveniles may disperse several hundred kilometres. The species is not considered to be seriously at risk to disturbance at most breeding localities. However, modern expansion of fishing activities may affect the very large populations of Ichaboe and Mercury Islands.     

Evaluating the Impact of Handling and Logger Attachment on Foraging Parameters and Physiology in Southern Rockhopper Penguins

Logger technology has revolutionised our knowledge of the behaviour and physiology of free-living animals but handling and logger attachments may have negative effects on the behaviour of the animals and their welfare. We studied southern rockhopper penguin (Eudyptes chrysocome) females during the guard stage in three consecutive breeding seasons (2008/09−2010/11) to evaluate the effects of handling and logger attachment on foraging trip duration, dive behaviour and physiological parameters. Smaller dive loggers (TDRs) were used in 2010/11 for comparison to larger GPS data loggers used in all three seasons and we included two categories of control birds: handled controls and PIT control birds that were previously marked with passive integrative transponders (PITs), but which had not been handled during this study. Increased foraging trip duration was only observed in GPS birds during 2010/11, the breeding season in which we also found GPS birds foraging further away from the colony and travelling longer distances. Compared to previous breeding seasons, 2010/11 may have been a period with less favourable environmental conditions, which would enhance the impact of logger attachments. A comparison between GPS and TDR birds showed a significant difference in dive depth frequencies with birds carrying larger GPS data loggers diving shallower. Mean and maximum dive depths were similar between GPS and TDR birds. We measured little impact of logger attachments on physiological parameters (corticosterone, protein, triglyceride levels and leucocyte counts). Overall, handling and short-term logger attachments (1–3 days) showed limited impact on the behaviour and physiology of the birds but care must be taken with the size of data loggers on diving seabirds. Increased drag may alter their diving behaviour substantially, thus constraining them in their ability to catch prey. Results obtained in this study indicate that data recorded may also not represent their normal dive behaviour.     

Group Size in Foraging African Penguins (Spheniscus demersus)

Diving synchrony was examined for varying group sizes of African penguins (Spheniscus demersus) travelling to their foraging grounds from their breeding islands. Groups of fewer than 12 birds always dived synchronously, whereas groups of more than 17 birds always dived asynchronously. Since travelling penguins do not dive deeply, large groups of birds can remain together irrespective of diving synchronization. Observations from boats showed that foraging penguins rarely occurred in groups of more than 17 birds. We suggest that groups of penguins that do not have synchronized dives cannot forage effectively, because foraging penguins dive deeply.     

5. FIELD OBSERVATIONS,NORTHERN PROVINCE,NORTHERN RHODESIA

Cooper, J. 1986. Biology of the Bank Cormorant, Part 4: Nest construction and characteristics. Ostrich 57: 170–179.

Bank Cormorants Phalacrocorax neglectus construct their nests of material gathered by diving. Males undertake diving bouts of approximately 3–5 min, made up of several dives lasting on average 28 s. Nest material is gathered throughout the breeding cycle: number of diving bouts per day varies from a mean of 7,6 during pre-egg laying to 1,5 bouts per day when rearing young in the nest. Nest building recommences within 24 h of loss of nest due to storms. Both sexes occasionally steal nest material from the nests of neighbours. Bank Cormorants sometimes defecate onto their nests. This is assumed to make the nests better able to withstand rough seas. Nest construction takes approximately 34 d, a period similar to that of other ground-nesting species of cormorants. Construction of a nest in 34 d represents 238 diving bouts of a total duration of 18 h. Nests are heavy (up to 6 kg) and are made up primarily of seaweed. Feathers, sticks and artificial material are also incorporated into the nest. Bank Cormorant nests are large (up to 54 litres in total volume). Nests do not change significantly in size between egg laying and hatching. Nests in which at least one egg hatches are larger in all dimensions measured than those in which no eggs hatch. Nests are larger at the time of laying of repeat clutches than at the time of laying the first clutch. The Bank Cormorant's seaweed nest has enabled it to breed on bare offshore rocks where no nest material exists. The species' large nest is a necessary prerequisite for successful breeding close to the sea.     

The use of supplementary foods by Australian magpies Gymnorhina tibicen: Implications for wildlife feeding in suburban environments

Abstract The provision of foods to wild birds is an extremely common practice among householders throughout the Western world. Nonetheless, concerns over potential impacts of the practice are currently being raised, including the possibility that some species may become reliant on human‐provided food. We compared the foraging and breeding ecology of pairs of fed and unfed Australian magpies Gymnorhina tibicen living in suburban environments in Brisbane, Queensland, Australia. Detailed behavioural observations of both foraging and chick provisioning were made for males and females of both groups throughout the breeding season. Natural foods dominated the diets of both fed and unfed magpies, making up 76% and 92% of all items consumed respectively. During the morning, fed magpies obtained fewer food items during ground foraging than did unfed birds, apparently because they visited suburban feeding stations more often during the early part of the day. At other times, the amount of food items obtained during foraging was similar for both fed and unfed birds. Magpies utilizing suburban feeding stations started all breeding activities significantly earlier than unfed magpies, except during the fledgling phase. Both fed and unfed magpie parents provisioned their chicks predominantly with natural food. Magpies were not reliant or dependent on supplementary food provided by wildlife feeders at any time during the breeding season. Although many magpies did utilize suburban feeding stations extensively, they continued to forage for and provision their chicks with natural food.     

Effects of prey abundance on the foraging behaviour, diving efficiency and time allocation of breeding Guillemots Uria aalge

The foraging behaviour of Guillemots Uria aalge at sea was compared between 2 years of radically different food abundance. Radio telemetry was used to determine foraging locations and diving patterns. In the poor compared with the good food year, foraging trips were much longer, the birds foraged more than six times further from their breeding sites, they spent over five times as much time diving when at sea and their estimated energy expenditure was twice as great. Time spent foraging in the poor food year was at the expense of time spent sitting at the colony. The duration of a foraging trip was a poor indicator of distance travelled but a good indicator of the amount of time spent diving. Mean dive durations, surface pause durations and interbout periods did not differ between years, but individuals made more than four times as many dives per diving bout in the poor food year. Surface pause lengths did not vary with water depth in either year. In the poor food year, birds made shorter surface pauses for a dive of a gi^ven duration than in the good food year, possibly accepting a lactic acid debt in order to maximize searching time, The duration of the interbout period was positively related to the number of dives in the previous bout, and dives tended to get shorter in long diving sequences, suggesting possible exhaustion effects. These data demonstrate that breeding Guillemots have the capacity to adjust their foraging behaviour and time budgets in response to changes in food abundance, but this flexibility was not sufficient to compensate fully for the very low food abundance experienced by birds in this study.     

Relationship between reversed sexual dimorphism,breeding investment and foraging ecology in a pelagic seabird,the masked booby

Reversed sexual dimorphism (RSD) may be related to different roles in breeding investment and/or foraging, but little information is available on foraging ecology. We studied the foraging behaviour and parental investment by male and female masked boobies, a species with RSD, by combining studies of foraging ecology using miniaturised activity and GPS data loggers of nest attendance, with an experimental study where flight costs were increased. Males attended the chick more often than females, but females provided more food to the chick than males. Males and females foraged during similar periods of the day, had similar prey types and sizes, diving depths, durations of foraging trips, foraging zones and ranges. Females spent a smaller proportion of the foraging trip sitting on the water and had higher diving rate than males, suggesting higher foraging effort by females. In females, trip duration correlated with mass at departure, suggesting a flexible investment through control by body mass. The experimental study showed that handicapped females and female partners of handicapped males lost mass compared to control birds, whereas there was no difference for males. These results indicate that the larger female is the main provisioner of the chick in the pair, and regulates breeding effort in relation to its own body mass, whereas males have a fixed investment. The different breeding investment between the sexes is associated with contrasting foraging strategies, but no clear niche differentiation was observed. The larger size of the females may be advantageous for provisioning the chick with large quantities of energy and for flexible breeding effort, while the smaller male invests in territory defence and nest guarding, a crucial task when breeding at high densities. In masked boobies, division of labour appears to be maximal during chick rearing—the most energy-demanding period—and may be related to evolution of RSD.     

Social context and prey composition are associated with calling behaviour in a diving seabird

Social cohesion and prey location in seabirds are largely enabled through visual and olfactory signals, but these behavioural aspects could potentially also be enhanced through acoustic transfer of information. Should this be the case, calling behaviour could be influenced by different social–ecological stimuli. African Penguins Spheniscus demersus were equipped with animal-borne video recorders to determine whether the frequency and types of calls emitted at sea were dependent on behavioural modes (commuting, sedentary and dive bout) and social status (solitary vs. group). For foraging dive bouts we assessed whether the timing and frequency of calls were significantly different in the presence of schooling prey vs. single fish. The probability of call events was significantly more likely for birds commuting early and late in the day (for solitary birds) and during dive bouts (for groups). During foraging dive bouts the frequency of calls was significantly greater for birds diving in the presence of schooling fish and birds called sooner after a catch in these foraging scenarios compared with when only single fish were encountered. Three call types were recorded, 'flat', 'modulated' and 'two-voice' calls, but there was no significant relationship detected with these call types and behavioural modes for solitary birds and birds in groups. The results of this study show that acoustic signalling by African Penguins at sea is used in a variety of behavioural contexts and that increased calling activity in the presence of more profitable prey could be of crucial importance to seabirds that benefit from group foraging.     

Seasonal and annual variation in the diving behaviour of Hutton's shearwater (Puffinus huttoni)

ABSTRACT

With the development and implementation of tracking technology, we are now able to monitor the foraging behaviour of seabirds while at sea. Time-Depth Recorders (TDRs) were fitted to Hutton's shearwaters (Puffinus huttoni), an endangered endemic New Zealand species, to measure how diving behaviour varies over the breeding cycle. Hutton's shearwaters (～350?g) dive up to 339 times per day (average 68.8) at depths to 35?m (average 5.6?m), and for periods up to 60?s (average 19.2?s). Incubating birds dived deeper than birds feeding chicks, and a significant difference in diving depth and dive duration were detected at different times of the day. Neither dive frequency nor dive duration differed significantly between years, but there was some annual variation in dive depths. The temporal variation we observed in the diving behaviour of Hutton's shearwaters suggests they are likely to exploit different types of pelagic prey at different stages in their breeding cycle. With on-going changes in the marine environment, monitoring changes in feeding behaviour using TDRs may provide a way to assess environmental change and improve the conservation of this species.     

Maternal attendance behaviour of sympatrically breeding Antarctic and subantarctic fur seals,Arctocephalus spp., at Macquarie Island

Maternal attendance behaviour was studied in Antarctic (Arctocephalus gazella) and subantarctic fur seals (Arctocephalus tropicalis) which breed sympatrically at subantarctic Macquarie Island. Data on attendance were obtained using telemetric methods. Both species undertook two types of foraging trips: overnight foraging trips which were of less than 1 day duration and occurred exclusively overnight, and extended foraging trips which lasted longer than 1 day. The mean duration of overnight foraging trips was 0.43 and 0.39 days, while the duration of extended foraging trips was 3.6 and 3.8 days in A. gazella and A. tropicalis, respectively. The duration of overnight and extended foraging trips did not differ significantly between species. Two types of shore attendance bouts that differed in duration were also observed in these species. Short attendance bouts lasted less than 0.9 days, while long attendance bouts lasted longer than 0.9 days. Short attendance bouts lasted 0.4 and 0.5 days, while long attendance bouts lasted 1.6 and 1.7 days in A. gazella and A. tropicalis, respectively, and did not differ significantly between species. The most significant differences between the attendance behaviour of both species was in the percentage of foraging time allocated to overnight foraging trips (15% and 25% in A. gazella and A. tropicalis, respectively), and the percentage of time spent ashore (30% and 38% in A. gazella and A. tropicalis, respectively). The nearness of pelagic waters to Macquarie Island is considered to be the main reason that lactating females are able to undertake overnight foraging trips. These trips may be used by females as a means of optimising the costs of fasting and nursing ashore. Females may be able to save energy by only nursing pups when milk transfer efficiencies are high, and reduce the time and energy costs of fasting ashore when milk transfer efficiency is low. Of the female A. gazella that still carried transmitters at the end of lactation, 83% continued regular attendance for between 21 and 150 days post-lactation (when data collection ceased). Overwintering of A. gazella females at breeding sites has not been previously reported in other populations. Accepted: 10 November 1998     

Foraging behaviour of chick-rearing rhinoceros auklets Cerorhinca monocerata at Teuri Island, Japan, determined by acceleration-depth recording micro data loggers

Flight and diving activity of rhinoceros auklets Cerorhinca monocerata breeding on Teuri Island, Japan, were monitored during the summers 1999 and 2000 using miniaturized time-depth and acceleration recorders. Birds made 14.5 dive bouts per day of on average 15.4 min duration, which consisted of on average 16.2 dives of 12.1 m depth and 42.7 s duration. Birds made 13.8±7.3 flight bouts per day, which lasted on average 11.5±4.5 min. Daily total flight duration was 2.7±1.7 h (range 54 s–5.1 h) and the mean potential foraging range was estimated to be 87 km (maximum 164 km). Most birds stayed at the colony or rested on the water surface during the night. Rhinoceros auklets dived more actively in early morning and in late afternoon than during mid-day. Compared to results from studies of time allocation in other alcids species, rhinoceros auklets spent longer time flying (3.3 hd⁻¹) and resting on water (13.1 hd⁻¹), and less time diving (3.1 hd⁻¹) and staying at the colony (4.4 hd⁻¹). These foraging patterns are probably related to the nest attendance pattern of rhinoceros auklets, i.e. leaving the colony early in the morning, staying at sea all the day and returning to the colony in the evening to provision their chicks.     

The reproductive tactics and activity patterns of solitary carnivores: the Iriomote cat

Felids are generally considered to be crepuscular and nocturnal in their activity, but few studies have attempted to analyze the variability of their activity patterns. We studied the daily activity of the Iriomote cat Prionailurus iriomotensis by radio-tracking on Iriomote Island, Japan. The general activity patterns of Iriomote cats showed slightly prevailing activity during dark periods of the day with particular peaks at dawn and dusk or during the early hours of the night. However, these patterns were clearly dependent on the sex and reproductive status of the cat. Peaks of cats’ activity coincided with those of their main prey. On average, the cats were active for thirteen hours per day. During the mating season, the rhythm of activity in males followed that of breeding females, but not that of non-breeding ones. Males exhibited 11% higher total daily activity and longer active bouts during the mating period than in the remainder of the year. Breeding females had additional mid-day activity peak during the nursing period, but their total time of activity per day was 16% lower than in the period of kittens’ independence. Their active bouts were shorter and more frequent during nursing than at other times. These results suggest that lactating females perform frequent movements to and from the den site to care for kittens. During the non-nursing period, females increased their activity, possibly in response to lowered prey abundance and the need of intensive foraging to recover after lactation. Seasonal and sexual variation of activity patterns in the Iriomote cats confirmed the existence of different reproductive strategies of males and females of these solitary carnivores.