Costs of reproduction in lizards: escape tactics and susceptibility to predation

Summary Increased basking and reduced agility in gravid female southern water skinks (Eulamprus tympanum) suggest that they will be more vulnerable to predators. However, gravid females shift their anti-predator tactics towards crypsis, by allowing potential predators (such as a human observer) to approach more closely than do males and non-gravid females. Gravid females were taken no more frequently than were non-gravid females or males when exposed to two types of natural predators, birds (kookaburra, Dacelo gigas) or snakes (common blacksnakes, Pseudechis porphyriacus) in field enclosures. Both these results suggest that the vulnerability of potential prey in this system is determined by the predator's probability of detecting a potential prey item, not its probability of capturing the prey item after detection. Hence, laboratory-based measures of performance demonstrating reduced escape speed may sometimes have little relevance to actual fitness under field conditions, if the probability that an animal will be taken by a predator depends primarily on whether or not it is seen, rather than on how quickly it can escape. Correspondence to: L. Schwarzkopf     

Does reproductive state affect a lizard's behavior toward predator chemical cues?

We examined how reproductive state affected shifts in behavior of adult female Podarcis sicula toward chemical cues from a natural snake predator (Coronella austriaca). The oviparous P. sicula lizards do not experience a major physical burden during reproduction, but gravid females substantially increase duration of basking. Therefore, gravid lizards are likely to experience a greater risk of mortality because they are exposed to predators for longer periods. Both gravid and non-gravid females shifted patterns of locomotion when confronted with snake chemical cues, but the change was notably larger when females were non-gravid. When non-gravid, lizards responded to predator scent by increasing the number of stand-ups and starts, while such a response was not observed when females were gravid. By contrast, gravid lizards clearly reduced the time spent basking in the presence of predator scent, whereas no change in basking behavior was observed when females were non-gravid. Thus, females exhibit differential behavioral responses to predator scents that is dependent on reproductive state.     

Life history of the nematode Theristus anoxybioticus from sublittoral muddy sediment at methane seepages in the northern Kattegat,Denmark

Field and laboratory studies from 1989 to 1994 of the nematode Theristus (Penzancia) anoxybioticus Jensen, 1995 inhabiting muddy sediment at methane seepages in the northern Kattegat, Denmark, have revealed a series of biological features of adults and juveniles not observed before in free-living marine nematodes. Reproductive adults were present in April and May only and inhabited the uppermost cm of sediment, while juveniles inhabited deep anoxic and sulphidic sediment layers during the remaining months of the year. This points to a downwards migration of newly hatched juveniles and an upwards migration during adulthood. Gut content analyses showed that adults feed mainly on benthic diatoms by swallowing whole cells while juveniles probably feed on bacteria; this differentiation in food clearly relates to the main bulk of potential food items in their respective microhabitats. The generation time is estimated to be one year, with a juvenile life span of about nine months, which is four to eight times longer than that found in related aerobic species of similar body size. Adults survived in oxic water but 50% died within 15 h in hypoxic water (5 to 6% atm sat), and all died within 12 h in anoxic water. Juveniles on the other hand died within 0.5 h in oxic water but survived anoxic water for at least 15 d. These results suggest that juveniles of T. anoxybioticus are adapted to an anaerobic life style.     

Kinship and social tolerance among female and juvenile wild house mice: kin bias but not kin discrimination

Kin-biased social tolerance among house mice has been interpreted in terms of kin discrimination. However, several lines of evidence suggest it may instead be an incidental artifact of group member discrimination. This leads to very different predictions about the social consequences of relatedness within and between social groups. Social interactions between wild-stock adult female and juvenile house mice (Mus domesticus) established in neighbouring territorial groups within enclosures reveal relatedness to dominant males within groups as the major factor determining social tolerance of juveniles by females. Relatedness to the female herself had no significant independent effect on responses indicating tolerance. Females were generally more aggressive toward neighbouring-group juveniles (all unrelated to females) compared with those from their own group (all related to females), but were most aggressive toward neighbouring juveniles sired by the neighbouring dominant male. They were also more aggressive toward their own-group juveniles that had been sired by the neighbouring dominant but only when encountered in the neighbouring territory and with a greater bias against female juveniles. Females were least aggressive toward own-group juveniles sired by their own-group dominant male. The sire-bias in tolerance among females is similar to that reported among the dominant males themselves in an earlier study. As a result of the combined sire-bias in tolerance by adult males and females, juveniles sired by their own-group dominant males become less likely to intrude into a neighbouring territory with time. Overall, the results suggest that differences in social tolerance reflect discrimination on the basis of social group membership rather than relatedness between interactants and thus provide strong experimental evidence in support of incidental kin bias rather than kin discrimination.     

Mating patterns and reproductive success in the bushy-tailed woodrat (Neotoma cinerea), as revealed by DNA fingerprinting

The mating patterns and reproductive success of the bushy-tailed woodrat (Neotoma cinerea) were investigated over a 3-year period (1992–1994) using DNA fingerprinting. Paternity was determined by genetic analysis of 58 juveniles of known maternity from 35 litters. Analysis of DNA fingerprints revealed that all offspring within a litter were fathered by a single male; the statistical probability of detecting multiple males mating with a female was high, indicating that multiple paternity would have been detected had it occurred. However, individual males did not father more than one litter from a given female either within or between years. At least 75% of females and 57% of males successfully produced offspring each year. The finding that all littermates are first-order relatives may contribute to the high level of female cooperation in this species. Received: 28 May 1997 / Accepted after revision: 22 March 1998     

The Arctic sea butterfly<Emphasis Type="Italic"> Limacina helicina</Emphasis>: lipids and life strategy

The sea butterfly Limacina helicina was collected from May to September 2001 in Kongsfjorden, Spitsbergen, to investigate population structure and body and lipid composition with regard to life cycle and reproductive strategy. Veligers and juveniles were only found in late autumn and spring, whereas females occurred from July to September. The size of the females increased until mid-August and decreased in September. Dry and lipid mass were closely related to size; dry mass increased exponentially and lipids linearly with size. The lipid content was highest in veligers (31.5% of dry mass) and juveniles (23.6%) but low in females (<10%). Phospholipids were the dominating lipid class followed by triacylglycerols. Females, veligers, and egg ribbons, all from September, were richest in phospholipids. Juveniles contained the highest amounts of triacylglycerols, whereas females had low levels in July and the beginning of August. In mid-August, levels of triacylglycerols were variable and higher. This suggests that females were in their main spawning period and the high variability in triacylglycerols points to different stages within the spawning cycle. Enhanced amounts of free fatty acids in females from July may be related to gonad development. The 16:1(n-7) fatty acid was more dominant in spring whereas 18:4(n-3) increased in summer and autumn, which reflects a change in diet from diatom-dominated food items in spring to dinoflagellates in summer/autumn. Small amounts of long-chain monounsaturated fatty acids suggest ingestion of copepods, and the fatty acid composition of veligers feeding on particulate matter. L. helicina has a one-year life cycle with peak spawning in August and over-winters as veligers that may grow to juveniles during the winter period. They metamorphose into juveniles during spring, develop to males in early summer, and mature into females in July and August.Communicated by M. Kühl, Helsingør     

Living in the front: Neomysis americana (Mysidacea) in the Río de la Plata estuary, Argentina-Uruguay

The Río de la Plata is one of the main estuarine systems of South America. It is characterized by a salt wedge regime, a well-developed bottom salinity front, and a maximum turbidity zone associated with it. We described, for the first time, the spatial distributional patterns of Neomysis americana, the most abundant mysid and the main food item for juvenile fishes in this estuary. We analyzed the link between mysid distribution and abundance and the bottom salinity gradient. A total of 242 plankton samples were taken from the Río de la Plata estuary in spring and fall between 1991 and 2001. Bottom salinity gradient was quantified from grids created on the basis of 348 oceanographic stations. The N. americana population was characterized by high abundances (up to 2500 ind. m⁻³), with juveniles, males, gravid and non-gravid females present in both spring and fall of different years. N. americana distribution followed the position of the bottom salinity front in different years and seasons. Pearson’s correlation analysis between mysid abundance and bottom salinity gradient confirmed the association of mysids with the bottom salinity front (maximum salinity gradient). No correlation was detected between mysid abundance and salinity per se or temperature (neither in spring nor in fall). We speculate that mysids concentrated at the front could take advantage of the high concentration of detrital material for feeding. The results of our work highlight the importance of the magnitude of salinity gradient for the ecological processes of a salt-wedge estuary like the Río de la Plata. The analysis of the spatial distribution of gradient values presented in this work also constitutes a useful tool to locate key ecological areas such as fronts.     

Distribution,growth and life cycle of the mesopelagic amphipod Primno abyssalis (Hyperiidea: Phrosinidae) in the southern Japan sea

The vertical distribution, growth, maturation, brood size and life cycle of the hyperiid amphipod Primno abyssalis (formerly P. macropa) were investigated using seasonal samples collected from Toyama Bay, southern Japan Sea, during the period June 1986 to September 1992. Over four different seasons of the year, P. abyssalis was most abundant in the 200 to 350 m strata at night and the 350 to 400 m strata during the day, indicating 100 to 150 m as the general distance of diel vertical movement. Some differences in vertical migrating behavior were noted among juveniles, adult males and females. Population-structure analysis revealed the occurrence of three cohorts aged 0+, 1+ and 2+ yr. Growth as body length in this species is linear with time. Estimated time to complete one life cycle is 1.8 to 2.5 yr for females, but only 0.8 yr for males. Maximum longevity is 2.8 yr. Instar analysis based on the segment number of pleopod rami indicated that newly hatched juveniles molt ten times to reach adult male, and four more times to reach adult female. Adult instar number was found to be only 1 for males and 5 for females. Ovigerous females occurred throughout the year, but the annual peak of release of juveniles from the female's marsupium is estimated to be arly March. Brood size was not correlated with female size, a maximum brood size of 214 eggs was recorded. The dry and ash-free dry weights of instars suggested that juveniles in the female marsupium, adult males, and older adult females are less active feeding or non-feeding stages. Except for the reduced growth rate and the occurrence of small, short-lived males, most characteristics of P. abyssalis are consistent with the present view of the life modes of mesopelagic animals, including linear growth in length, aseasonal reproduction, and smaller brood size coupled with larger eggs.     

The ecophysiological complex of Bathyporeia pilosa and B. pelagica (Crustacea: Amphipoda). II. Effects of exposure

The resistance to high and low temperatures, starvation and desiccation effects in the sand dwelling amphipods Bathyporeia pilosa Lindström and B. pelagica (Bate) have been tested. B. pilosa was the more tolerant species in all experiments. Gravid females showed a greater resistance to high temperatures than adult males in both species, and starvation tolerance was in the order gravid females>juvenile males> adult males. Time-temperature relationships, however, would seem to be of greater ecological significance than upper or lower lethal temperatures, and desiccation effects become obvious well within any limits set by starvation. Exposure would appear to be an important concept limiting the intertidal distribution of sand dwelling animals, particularly those confined to the surface layers of sediment. The amplitude, rate and degree of environmental change and its effect on feeding and reproduction offer severe limitations to the distribution of the two Bathyporeia species studied.     

Flight initiation distance decreases during social activity in lizards (<Emphasis Type="Italic">Sceloporus virgatus</Emphasis>)

Flight initiation distance, the predator–prey distance when escape begins, is predicted by escape theory to decrease if fleeing entails loss of benefits. Shortening of flight initiation distance during social interactions is known only in males and only in a few species. In a previous study, male, but not female, Sceloporus virgatus lizards had shorter flight initiation distance when interacting with tethered conspecifics. Females in that study were not gravid or close to ovulating. I predicted that flight initiation distance would be shorter in gravid females that perform sidle-hopping displays to reject courtship than in lone females. I tested this prediction and examined effects of social interactions by males with free-ranging conspecifics to ensure that previous findings were not artifacts of tethering and experimental introduction of conspecifics. Flight initiation distance was shorter in females when interacting with males than when alone; it was also shorter in males interacting with either sex. Thus, when beneficial for reproductive reasons, social interaction affects flight initiation distance in females, but at other times, it does not. Lesser shortening of flight initiation distance in females than males may be a consequence of greater social benefit to males and protection of reproductive investment by females.     

Accumulation and release of fuel-oil-derived diaromatic hydrocarbons by the polychaete Neanthes arenaceodentata

Male and gravid female Neanthes arenaceodentata (Moore) (Polychaeta: Annelida) were experimentally exposed to a sublethal concentration of No. 2 fuel oil seawater extract for 24 h. Within 1 h both sexes had incorporated, in equal proportions, most of the diaromatic hydrocarbons (naphthalene, methylnaphthalenes) eventually accumulated. No net loss of accumulated naphthalenes was observed during the exposure period. When returned to hydrocarbon-free seawater, male worms slowly released naphthalenes down to undetectable (< 0.1 ppm) levels in 400 h (17 days). Gravid females retained essentially all of the originally accumulated naphthalenes for 3 weeks, at which time they released their eggs during fertilization, with subsequent dramatic decreases in naphthalenes content. Zygotes and trochophore larvae from exposed females contained 18 ppm naphthalenes, accounting for a majority of those amounts initially present in parent females. As trochophore larvae developed into 18-segment juveniles, naphthalenes concentrations dropped to near undetectable levels (0.4 ppm) in 21-day old juveniles (= 18-segment worms). When sampled at the next stage of development (32-segment worms) 5 days later (1050 h after exposure of gravid females), offspring were free of naphthalenes. Results suggest the importance of lipid-rich eggs in the accumulation and retention of diaromatic hydrocarbons by marine polychaetes.     

Prolonged copulation: A male ‘postcopulatory’ strategy in a promiscuous species,Lygaeus equestris (Heteroptera: Lygaeidae)

Summary Copulation in Lygaeus equestris L. (Heteroptera, Lygaeidae) is known to last 0.5–24 h. Variations in copula duration of field-collected insects were studied in the laboratory, and different hypotheses concerning the significance of prolonged copulations were tested.Through reciprocal matings with normal and sterile (irradiated) males, sperm displacement was estimated at about 90%. A male could thus increase his fitness by preventing subsequent matings of an inseminated female.Copulations with virgin insects, observed over 15 h, were classified in two categories: short (0.5–8.0 h) and long (> 15 h) duration (Fig. 1). No difference in the number of fertilized eggs was found between long and short copulations, and the insemination rate was highest during the first hour of a couplation (Fig. 2). Long-lasting couplations did not give rise to increased sperm displacement (Table 2). These results indicate that there is no difference in the amount of sperm transferred during short and long copulations and that no insemination takes place during the latter part of a prolonged copulation.Longer lasting copulations occurred when the sex ratio was male-biased than when it was female-biased (Fig. 1). The frequency of prolonged copulations was higher when the female was gravid than when she contained no eggs (virgin) (Figs. 3 and 4). Support is thus given to the hypothesis that prolonged copulation is a male postinsemination strategy to prevent subsequent matings of a female.Females subjected to male competition over a longer period, laid fewer eggs and died earlier than females that were mated but subsequently isolated from males (Fig. 5). Egg batches from females living with males were larger than batches from mated, isolated females (Table 3). This is probably due to a combination of many matings with short intermissions and prolonged copulations, both of which postpone oviposition.Comparisons between copulations of males with either gravid or virgin females during these experiments, led to the conclusion that short copulations with virgin females result from a male decision to terminate copulation after insemination is completed, whereas copula duration with gravid females is more likely to depend on a combination of male and female behavior.     

Harpacticoid copepod respiration: Enhydrosoma propinquum and Longipedia helgolandica

Oxygen consumption rates of 2 harpacticoid copepods, one benthic, Enhydrosoma propinquum (Brady) and one benthopelagic, Longipedia helgolandica (Klie), were determined and compared with those of the pelagic harpacticoid Euterpina acutifrons. The respiratory rates of the 3 species differed significantly and could be correlated with activity patterns and habitat preferences. E. propinquum consumed the least oxygen, E. acutifrons an intermediate amount, and L. helgolandica the most per unit dry weight. Gravid L. helgolandica had a lower metabolic rate than the non-gravid females. The lower values for the gravid females most likely reflected the metabolic activity of the developing egg mass. Harpacticoids may be important contributors to total meiobenthic metabolism, since they respire more per unit weight than do nematodes.Supported by NSF Grant GB-8189 to Duke University Cooperative Program in Biological Oceanography (BCC) and NSF Grant 7435 (WBV).     

Chemosensory mate recognition may facilitate prolonged mate guarding by male snow skinks, Niveoscincus microlepidotus

We provide evidence that male lizards can use chemosensory cues to identify individual females and probably therefore maintain long-term associations with these females in the wild. In the laboratory, males preferentially followed the scent trail of their vitellogenic female “partner” rather than that of another vitellogenic female. Our 5-year field study of the small viviparous scincid lizard (Niveoscincus microlepidotus) in alpine Tasmania showed that sexually mature males and females commonly formed “pairs” for long periods (on average 29 days). These pairs occurred primarily during the mating season, always involved one adult male and one adult female, and usually involved vitellogenic rather than gravid females. Our laboratory experiments suggest that a significant factor in maintaining those prolonged partnerships is male scent trailing of partners. Received: 28 October 1997 / Accepted after revision: 28 May 1998     

Indicators of physiological and immunological status of <Emphasis Type="Italic">Litopenaeus setiferus</Emphasis> wild populations (Crustacea,Penaeidae)

Constant environmental changes in oceanic and estuarine systems due to human activity in coastal zones require different tools and strategies for their study to be able to assess the health status of aquatic ecosystems. The Gulf of Mexico has a tremendous ecological importance because of its biological diversity. Baseline levels of blood metabolites and immune components were defined for wild tropical populations of Litopenaeus setiferus, an important shrimp species of the Gulf of Mexico. Osmotic pressure, oxy hemocyanin (OxyHc), glucose, proteins, cholesterol, lactate, and triacylglycerols (TAG) were used as indicators of physiological status; pro phenoloxydase (proPO) and phenoloxydase (PO) quantification of hemocytes, as well as respiratory burst, were used to assess the immunological status. Significant differences were observed in live weight of juveniles, males, and females of L. setiferus with mean values of 10.54, 35.05, and 40.33 g, respectively. A value of 460 mOsm/kg was recorded in the water where juveniles were sampled and 1,000 mOsm/kg in adult sampling zones. Consequently, the osmotic pressure of L. setiferus juveniles (718.02 mOsm/kg) was 22% lower than in adults (925 mOsm/kg); there were no differences between sexes. A significantly lower value in juvenile OxyHc was observed. This value was 24 and 16% lower than that observed in males and females, respectively. Hemolymph glucose, cholesterol, and triacylglycerol levels were not statistically different between shrimp stages, showing median values of 0.19, 0.32, and 0.39 mg/ml of glucose, cholesterol, and triacylglycerols, respectively. Hemocyanin was distributed normally in the juvenile–male population but not in females. A proportion between 52 to 81% of OxyHc of the total protein (OxyHc/Prot) was obtained for all stages. Digestive gland glycogen showed no normal distribution in juveniles and adults. Digestive gland glycogen was significantly higher in juveniles (median value of 2.64 mg/g) than in adults (median value of 1.58 mg/g). A higher value of lactate (0.11 mg/ml) was observed in juveniles than in males (0.09 mg/ml) and females (0.04 mg/ml). Granular cells (GC) corresponded to 36, 44, and 39% of the total hemocytes (TH) observed in juveniles, males, and females. ProPO showed no normal distribution in all stages. A high proPO activity was recorded in L. setiferus females that was 95% higher than in the juvenile–male group. Respiratory burst was divided into two groups, juveniles–males and females, and was distributed normally in both. Female respiratory burst values were 40 and 45% higher than those observed in the juvenile and male groups, respectively. There was an increment in OxyHc, proteins, PO, hemocytes, and respiratory burst with an increase in body weight. In contrast, the relationship of live weight to glucose, TAG, and proPO revealed no significant regressions. A significant decrease in cholesterol, glycogen, and lactate with an increase in body weight was also observed. We now have indicators of nutritional and immune status of juveniles and adults that can reveal changes in trophic relations and health at the population level. This will serve as a basis for using L. setiferus as environmental sentinels, as non-controversial and inexpensive models to evaluate the potential changes that human activities could exert on this important marine ecosystem.Communicated by P.W. Sammarco, Chauvin     

Social monogamy and extra-pair fertilization in an Australian lizard, Tiliqua rugosa

This study investigates social monogamy in the Australian sleepy lizard, Tiliqua rugosa. At a 70-ha site near Mount Mary, South Australia, we radio tracked 55 adult female and 39 adult male lizards during their spring activity periods. Each lizard was observed in 1–5 years. Females were observed with a single male partner on an average of 10.8 days per year, although in 17.3% of cases, females were observed on 2 or fewer days with a male. The most intense pairing period each year was 15 September–15 November when females were with male partners on an average of 36% of observation days. Partnerships lasted an average of 43.3 days each year. After mating in early November, the pairs separated. Observations of females pairing with other males were rare. Most males (82%) were also consistently monogamous, although 7 were observed pairing with 2 females within one season. To investigate paternity, we allowed 21 gravid females to give birth to 42 offspring in the laboratory. We determined genotypes at five polymorphic microsatellite DNA loci for the females, their male partners and their offspring. Four litters (19%) and 6 of the offspring from those litters (14.3%) showed evidence of extra-pair fertilization (EPF). Although the sample sizes are small, females of polygynous males were more likely to experience EPF. Received: 22 February 1998 / Accepted after revision: 23 May 1998     

The effects of size-dependent predation risk on the interaction between behavioral and life history traits in a stream-dwelling isopod

In field surveys, laboratory observations and field-based assays of behavior, I examined the effects of size-dependent predation risk on the interaction between size at reproductive maturity and maternal care behavior in the stream-dwelling isopod, Lirceus fontinalis. L. fontinalis exhibit population-specific sizes at reproductive maturity which result in population differences in predation risk during the adult phase. Females from streams containing salamander larvae (that prefer small prey) mature at large sizes and then become relatively safe from predation. Females from streams containing fish (that consume all size classes of prey equally) mature at small sizes and remain at risk. I tested whether these differences in expected survival were reflected in the behavior of females during the maternal phase (i.e., the period during which females exhibit maternal care). Female L. fontinalis carry developing juveniles inside a brood pouch. I simulated predatory attacks on gravid female L. fontinalis from the different population types and found that female behavior correlated with population differences in risk. When “attacked”, females from streams with predatory fish (that experience high risk to adult females) released juveniles from the brood pouch, whereas females from populations with predatory salamander larvae (that pose relatively little risk to adult females) did not release juveniles. I discuss the results with reference to the joint evolution of behavioral and life history traits. Received: 6 March 1996 /Accepted after revision: 12 August 1996     

Sexual selection favours small and symmetric males in the polygynous greater sac-winged bat Saccopteryx bilineata (Emballonuridae, Chiroptera)

We investigated how morphological traits of territorial males in the polygynous bat Saccopteryx bilineata were related to their reproductive success. Because of the frequency of aerial courtship displays and defence manoeuvres, and the high energetic costs of flight, we expected small and symmetric males to be better able to court females on the wing and to monopolize copulations with females in their harems. We predicted that small and symmetric males would sire more offspring within the colony and a larger portion of the young born within their harem than large or asymmetric males. We measured size and fluctuating asymmetry of 21 territorial males and analysed their reproductive success in 6 offspring cohorts (n=209 juveniles) using 11 microsatellite loci. As predicted, small and symmetric males had, on average, a higher reproductive success in the colony than large and asymmetric males. The percentage of young sired by males within their harem increased as males decreased in size, but was not influenced by fluctuating asymmetry. As fluctuating asymmetry of males correlated with their reproductive success within the colony but not within their harems, we infer that fluctuating asymmetry is probably related to female choice, whereas male size is probably important for harem defence on the wing.Communicated by G. Wilkinson     

Temporal and vertical distribution of two ecologically different calanoid copepods (<Emphasis Type="Italic">Calanoides carinatu</Emphasis>s Krøyer 1849 and <Emphasis Type="Italic">Lucicutia grandis</Emphasis> Giesbrecht 1895) in the deep waters of the central Arabian Sea

The full-depth vertical distributions of males, females and juveniles of Lucicutia grandis and Calanoides carinatus are presented from the surface down to depths of 3,900 m in the central Arabian Sea at the end of the NE monsoon and the February and October inter-monsoonal periods. L. grandis is a steady component and C. carinatus is a temporal resident of the central Arabian Sea oxygen minimum zone. Maximum abundances of C. carinatus and L. grandis juveniles were found at depths around 600 m, whereas the center of the distribution of L. grandis adults was somewhat deeper at 950 m. Both copepods are indicator species which can help to understand the functioning of the mesopelagic Arabian Sea ecosystem.     

Dominance behaviour,body weight and fat variations,and partial migration in European blackbirds Turdus merula

Summary A partially migratory population of European blackbirds Turdus merula was studied in central Europe during the non-breeding season. Body weight and the amount of subcutaneous fat deposits showed seasonal variations typical for each age and sex category. Juveniles were fattest in autumn and spring and both juveniles and females lost fat and body weight in mid-winter. Adults and males reached their maximum body weight in mid-winter. These findings together with data from Schwabl (1983) suggest that juveniles and females constitute the main fraction of those birds migrating from the area in autumn. In winter mainly adults and males have access to food resources. These patterns may be regulated by dominance behaviour, since adults and males are usually dominant over juveniles and females throughout the non-breeding season. Subordinate birds that stay in the area probably suffer from a higher winter mortality but have the advantage of early breeding.