Sibling recognition in Richardson's ground squirrels (Spermophilus richardsonii)

Summary Young from 10 litters of Richardson's ground squirrels were crossfostered within 24 h of birth. Litters, composed then of uterine-sibs and fostered non-sibs, were caged together until 37 days of age (1 week after weaning), at which time the juveniles were separated and caged individually. At a mean age of 110 d squirrels were tested for recognition during 10-min trials in an arena using the following pair-types: sibs reared together (SRT), sibs reared apart (SRA), non-sibs reared together (NSRT), and non-sibs reared apart (NSRA). Juvenile S. richardsonii displayed an ability, determined prenatally or soon after birth, to recognize their biological siblings. Familiarity, based upon association before and at the time of weaning, also affected discrimination between conspecifics, although this effect was less apparent.     

Alarm calling behavior of the thirteen-lined ground squirrel,Spermophilus tridecemlineatus

Summary Alarm calling in a population of thirteenlined ground squirrels, Spermophilus tridecemlineatus, was studied over a three-year period. Data on ground squirrel reactions to human and canine approaches and to the approach or presence of avian predators were used to quantify alarm calling behavior.The results support the hypothesis that alarm calling in this species functions to warn genetic relatives. Human and canine approach-elicited calls were most frequently given by mothers and their recently emerged young; adult males and nonparous females rarely called. The onset of maternal calling coincided with the aboveground appearance of a mother's own litter, and both juvenile and maternal calling were terminated at approximately three weeks post-emergence. Alarm calls were rarely emitted during encounters with avian predators.Alarm calling behavior in S. tridecemlineatus thus appeared to be dependent upon the presence of newly emerged juvenile relatives. To investigate whether the population structure of S. tridecemlineatus was perhaps incompatible with the evolution of alarm calling directed toward adult relatives, the distance between the home ranges of adult relatives and the distance over which the alarm vocalization is audible to ground squirrels were measured. The results revealed that females were likely to have adult relatives relatives residing within audible range of the call.     

Mating competition in an asocial ground squirrel,Spermophilus tridecemlineatus

Summary Male thirteen-lined ground squirrels (Spermophilus tridecemlineatus) are subject to three forms of intrasexual competition: competition over location of spatially scattered females, overt conflict over access to those females, and sperm competition. The likelihood of both overt conflict and sperm competition varies between seasons; changes in male density, female density, and breeding synchrony were investigated as possible sources of that variability. Male density was artificially reduced mid-way through one mating season. The results of that experiment, as well as subsequent between-season comparisons, indicate that fluctuations of male density have hardly any effect on levels of overt conflict and sperm competition. By contrast, those forms of intrasexual competition are influenced strongly by female density and/or breeding synchrony.     

Développement larvaire de deux Tellinacea,Scrobicularia plana (Semelidae) et Donax vittatus (Donacidae)

Scrobicularia plana Da Costa and Donax vittatus L. were reared in the laboratory through settlement. Fertilizable eggs were obtained by perfusing the ovary with 5% 0.1 M ammonium hydroxyde in sea water. S. plana eggs have a thick chorion, inside which the early larval stages develop; they hatch as straight-hinge larvae more than 60 h after fertilization. This brood protection is considered to be an adaptation to osmotic pressure changes and pollution in the environment. D. vittatus eggs have a very thin chorion and are unprotected. Further development is planktotrophic and very similar for the two species. Under laboratory conditions, the pediveliger stage is attained 3 weeks after fertilization and settlement occurs 1 week thereafter. S. plana spat stop growing until a suitable substratum is available. Meanwhile they undergo a byssus drifting important for postlarval dispersion. Post larvae resume growth as soon as a small quantity of fine sand is added to the rearing jar. The exhalant siphon is developed first, when the post larvae reach a length of 600 m; the inhalant siphon is formed later, at a length of approximately 900 m. Evolution from the larval hinge to the juvenile hinge stage occurs sooner in S. plana than in D. vittatus. Comparison of laboratory larval development with field development indicates that spawning occurs in June and August for S. plana in North Wales (UK).     

Kinship,demography, and belding's ground squirrel nepotism

Summary Social behavior and demography of a freeliving population of individually marked Belding's ground squirrels (Spermophilus beldingi) were studied from 1974 through 1980 at Tioga Pass, California. Relative frequencies of fighting, chasing, cooperation in chasing conspecifics, and assisting conspecifics who were being chased were recorded for adult (1-year-old) female kin of nine different degrees of matrilineal relatedness. The animals' mortality and dispersal patterns were also analyzed.Mothers and daughters, littermate sisters, and nonlittermate (half-) sisters were cooperative; their cooperation varied in proportion to relatedness. In contrast, grandmothers and granddaughters, aunts and nieces, great grandmothers and great granddaughters, aunts and half-nieces, first cousins, and first cousins once removed did not cooperate; their behavior was indistinguishable from that of nonrelatives.While there were no consistent spatial differences among the burrows of the nine categories of female kin, only mothers and daughters, littermate sisters, and nonlittermate sisters were consistently alive simultaneously. The apparent correspondence between the relatives that received social favoritism and those that consistently co-occurred suggests that demography, particularly survivorship, may have determined the extent of ground squirrel favoritism, while kinship influenced its pattern.     

Responses of nymphs of desert locust,Schistocerca gregaria to volatiles of plants used as rearing diet

Summary Gregarious nymphs of the desert locust,Schistocerca gregaria (Forsk.) (Orthoptera: Acrididae) were more attracted to volatiles from mechanically damaged food plants used for rearing than to either the undamaged or damaged food plants not used as diet in Y-tube olfactometer assays. Comparative analysis of the volatile emissions from plants used for rearing and food plants not used for rearing,e.g. Sorghum bicolor, Pennisetum clandestinum, Schouwia thebaica, wheat (Triticum sp., var. Nyangumi),Zygophyllum simplex, Heliotropium undulatum andTribulus terrestris was carried out by GC, GC-EAD and GC-MS. Significant quantitative and qualitative differences were found in the volatile emissions and olfactory responses of nymphs in GC-EAD assays. Up to 33 compounds were identified in volatiles of the plants of which 9 evoked EAGs. EAG-active components included common green leaf compounds (E)-2-pentenal, (E)-2-hexenal, 4-methyl-3-pentenal, (E)-3-hexenyl acetate, (Z)-3-hexenyl acetate, (Z)-2-hexenyl acetate, (Z)-3-hexen-1-ol and (Z)-2-hexen-1-ol. (Z)-3-Hexenyl butyrate and (Z)-3-hexenyl isovalerate were detected in stimulatory amounts only in the volatiles ofS. thebaica. (E, Z)-2,6-Nonadienal was detected as a component in the volatiles ofT. terrestris and was highly stimulatory. In EAG assays with seven common green leaf volatiles, (Z)-3-hexenyl acetate was most stimulatory while hexanal was the least. No significant differences were recorded between antennal responses of males and females to the tested compounds. These results are discussed with regard to current hypotheses on host plant recognition through detection of their airborne volatiles and the learning behaviour by nymphs ofS. gregaria.     

The culture and maintenance of the loliginid squids Sepioteuthis sepioidea and Doryteuthis plei

A technique for rearing the loliginid squids Sepioteuthis sepioidea and Doryteuthis plei is reported. Specimens of the former were reared from eggs to sexually mature adults, and maintained for a maximum of 146 days; adult D. plei were maintained for 38 days. Choice and quantity of food was most important for the survival of all sizes, particularly young squid. Newly hatched specimens thrived on Mysidium columbiae. Both species fed at a rate of 30 to 60% of their body weight daily; starvation occurred when intake fell below 10 to 15%. Food conversion efficiency averaged between 10 to 20%. Growth was rapid and steady. S. sepioidea grew to a maximum of 105 mm and 77 g in less than 5 months; D. plei grew an estimated 20 mm/month. Experimental data indicate a lethal minimum salinity for both species at about 27. Lethal minimum and maximum temperatures for young S. sepioidea are 17.5° to 18.0°C, and 32.5° to 33.0°C, respectively. Young consume 0.64l O₂/mg wet weight/h. Opaque tanks, with a semi-natural bottom substrate and special ultra-violet (UV) illumination, are advantageous for rearing and maintenance.Contribution No. 1305 from the Rosenstiel School of Marine and Atmospheric Science, University of Miami, USA. This work was supported in part by the National Science Foundation Grant GB-5729 X. This paper constitutes part of a doctoral dissertation on squid rearing, maintenance and behavior. Aspects of the behavioral studies will be published elsewhere.     

Intra-colony relatedness and sociality in the allodapine bee Exoneura bicolor

Summary Overwintered and newly co-founded nests of Exoneura bicolor exhibit different grades of sociality. Intra-colony relatedness was estimated for adults and female brood in both nest types using allozyme data and a multiallelic estimator. The higher relatedness among female brood from overwintered nests compared to newly-founded nests is consistent with the difference in sociality between these colony types (semisocial versus quasisocial). However, intra-colony relatedness among adults is higher in cofounded nests than in overwintered nests, suggesting that differences in sociality are determined by benefit/cost ratios associated with non-reproductive strategies, rather than relatedness between interactants. Rapid egg-production in newly founded nests allows eggs to be stockpiled. This reduceds the tasks available to non-reproductives in early phases of colony development and limits the payoffs available for reproductive altruism. It is suggested that the social flexibility characteristic of allodapines is a consequence of communal progressive rearing which allows many benefits from cooperative nesting to be gained without worker sterility.     

Binary mixtures of feeding deterrents mitigate the decrease in feeding deterrent response to antifeedants following prolonged exposure in the cabbage looper, <Emphasis Type="Italic">Trichoplusia ni</Emphasis> (Lepidoptera: Noctuidae)

Summary. The effect of rearing larvae of Trichoplusia ni on individual feeding deterrents or on binary mixtures of deterrents on their subsequent gustatory sensitivity was measured in paired choice leaf disc bioassays. Our working hypothesis was that mixtures of antifeedants (pure allelochemicals) would mitigate decreased feeding deterrent response following prolonged exposure in this generalist herbivore. Neonate larvae were reared on cabbage leaves treated with individual feeding deterrents (digitoxin, thymol, toosendanin or xanthotoxin), or with binary mixtures of these until the third instar. Feeding deterrent responses to each antifeedant or mixture was then determined in a leaf disc choice bioassay. All of the mixtures produced additive deterrence when presented to naïve larvae. Larvae reared on individual antifeedants showed a significantly decreased feeding deterrent response (except to digitoxin), whereas larvae reared on binary mixtures of antifeedants did not show a decreased feeding deterrent response to any of them. Such mixtures were synergistic in terms of their feeding deterrence to experienced larvae. Our experiment supports the hypothesis (Jermy 1986) that mixtures of deterrents can prevent decreased feeding deterrent response following prolonged exposure, and provides one explanation for the multiplicity of chemical defenses found in many plants.     

Kin preference behavior in Bufo boreas tadpoles

Summary The ontogeny of sibling recognition behavior was studied in the laboratory in tadpoles of the western toad (Bufo boreas boreas) to test the hypothesis that Bufo tadpoles associate with siblings and to compare this behavior with two species of anurans previously studied. Tadpoles reared exclusively with sibs demonstrated a preference to associate with sibs over non-sibs both early and late in development but tadpoles reared with sibs and non-sibs (mixed rearing groups) exhibited no preference. Larvae that developed a preference for sibs after being reared with them for 75 days lost this preference following exposure to a mixed group for 2 to 6 days. Additionally, larvae reared in a mixed group did not develop a preference for the familiar mixed group nor was a preference exhibited by individual larvae that were reared exclusively with 5 non-sibs. These results suggest that larvae learn by prior association to discriminate sibs from non-sibs and that preferences are rapidly modifiable following exposure to non-sibs. However, the latter two experiments suggest that social preferences are not totally labile and are not based entirely on familiarity. Field observations of larval activity, aggregation behavior, and dispersal patterns in B. boreas indicate that tadpoles mix with non-sibs from early larval stages throughout development. Because even short term exposure to nonsibs resulted in a loss of sib preference in our experiments, we suggest that a kin selection interpretation of B. boreas aggregation behavior is not parsimonious. Results differ with those of other studies of sib recognition in larval anurans.     

Photosynthesis-irradiance responses and photosynthetic periodicity in the sea anemone Aiptasia pulchella and its zooxanthellae

Sea anemones (Aiptasia pulchella) containing zooxanthellae (Symbiodinium microadriaticum) were maintained in a long-term laboratory culture on a 12 h light (100 E m^-2 s^-1):12 h dark cycle. Photosynthetic oxygen production was measured for the symbiotic association and for freshlyisolated zooxanthellae. Light utilization efficiencies () were similar for both sets of zooxanthellae, suggesting negligible shading of zooxanthellae by animal tissue in this association. Whereas freshly-isolated zooxanthellae were photoinhibited at high irradiances (800 to 1 800 E m^-2 s^-1), zooxanthellae in the host continued to function at photosynthetic capacity. Time of day may influence photosynthetic measurements in symbiotic organisms, as it was found that photosynthesis in A. pulchella followed a diel periodicity at both light-saturating (1 200 E m^-2 s^-1) and subsaturating (150 E m^-2 s^-1) irradiances. There was a peak period of photosynthesis between 12.00 and 14.00 hrs. Light stimulated dark respiration rates of A. pulchella. Dark respiration of sea anemones increased somewhat towards the end of the light cycle and was always greater after exposure to high irradiances.     

Laboratory rearing of the clupeid fish Harengula pensacolae from fertilized eggs

Pelagic eggs of the scaled sardine Harengula pensacolae (Goode and Bean), have been hatched and reared in the laboratory for the first time. Larvae were reared in two 75 l aquaria under constant illumination, at an average temperature of 26.2°C. Zooplankton collected in a 35 mesh net was fed to the newly hatched larvae, and the diet was supplemented later with Artemia salina nauplii and a pelleted food. Larvae hatched at 4 mm TL (total length), and metamorphosed about 25 days later at 25 to 30 mm TL. Survivors averaged 76 mm TL 100 days after hatching. Of the 500 incubated eggs, 2.8% survived until 20 days, after which no significant natural mortality occurred. Sources of natural mortality included starvation, a copepod parasite (Caligus sp.), and injuries from contact with the sides of the tank. Larvae began feeding at 4.5 mm TL on copepod nauplii averaging 62 in body width. Scaled sardines were photopositive throughout the larval stage.Contribution No. 149, Bureau of Commercial Fisheries Tropical Atlantic Biological Laboratory, Miami, Florida 33149, USA.     

Distribution of the early-life phase of the Antarctic squid Galiteuthis glacialis in relation to the hydrology of the Southern Ocean in the sector 15° to 30°E

During R.R.S. Discovery Cruise 100 in the Southern Ocean (1979), 620 specimens of the Antarctic cranchiid squid Galiteuthis glacialis (Chun) were caught in the RMT8 opening-closing net, to the south of the Antarctic Polar Front (APF). The catch was dominated by individuals <20 mm dorsal mantle length (DML) and, apart from two specimens, the rest of the catch was <60 mm DML. Two larger squid (255 mm DML) were both immature and apparently female. The species is concentrated at a depth of 300 to 400 m; there is evidence of ontogenetic spreading and a shift in the modal depth of the population from the 300 to 400 m horizon in daylight to the 200 to 300 m horizon in darkness. The collection confirms the circumpolar distribution of the species and supports previous evidence that it rarely, if ever, occurs to the north of the APF. As in Mesonychoteuthis hamiltoni, the only other species of cranchiid squid known to occur south of the APF, the early-life phase of G. glacialis is concentrated in the upper zone of the warm deep water beneath the Antarctic surface layer where, putatively, there is a zone of enhanced biological activity.     

Co-operative breeding by the Australian Bell Miner Manorina melanophrys Latham: A test of kin selection theory

Summary A population of Bell Miners Manorina melanophrys cosisted of two social units (clans), each consisting of one or more monogamous paris and numerous helpers. Individuals of a breeding pair occupied one home range, while non-breeding individuals occupied partially overlapping home ranges. Attendants (i.e. parents and helpers) showed clear preferences as to whom they aided (fed) and how much aid they provided. A significant correlation was found between the amount of aid given by an attendant and the genetic relationship between the attendant and the recipient. This provides strong quantitative evidence for the theory of kin selection. Only 14% of eggs laid produced juveniles which lived longer than 3 months. It is suggested that the extremely high juvenile mortality favours individuals which help relatives, even while such individuals have dependent offspring of their own.     

Mass culture of the rotifer Brachionus plicatilis and its evaluation as a food for larval anchovies

Growth rates of anchovy larvae, Engraulis mordax, reared for 19 days under constant environmental conditions on a diet of laboratory-cultured organisms, exceeded the growth rates of anchovies fed on a diet of wild plankton. The rotifer Brachionus plicatilis was found to be a nutritous food source when fed to the larvae in concentrations of 10 to 20/ml and in combination with the dinoflagellate Gymnodinium splendens (100/ml). Optimum conditions were determined for mass culture of the rotifer. A high food concentration was the most important parameter needed to assure a high yield of rotifers. Large volumes (464 I) of the unicellular flagellate Dunaliella sp. were cultured for feeding the rotifers. The rotifer culture technique described produces approximately 2.5×10⁶ organisms/day, providing a reliable food source for rearing studies. The lengths of B. plicatilis (without eggs) ranged between 99 and 281 , most rotifers being larger than 164 and less than 231 . Individuals weighed 0.16 g and contained 8×10^-4 cal.     

Bathymetric distribution of chaetognaths in the South Eastern Pacific Ocean

The bathymetric distribution of chaetognath fauna observed at planktonic stations in the South Eastern Pacific Ocean during three different expeditions, is presented quantitatively. The material from the cruises IFOP-01 (October–December, 1964) and IFOP-04 (November–December, 1965) was collected by vertical closing net, at regular intervals, from 2000 and 1000 m depth, respectively; the R.V. A. Bruun Cruise 13 data include vertical samples at usual depth intervals from 3000 m (see Table 12), Isaac-Kidd Midwater Trawl (IKMT) and surface collections. They allowed us to identify, for the first time, the meso and bathypelagic species of this region and to extend the longitudinal distribution of the epipelagic species, which had been previously limited to the coastal areas. Among the epiplanktonic species, it was shown that two, Sagitta serratodentata and S. bipunctata, do not penetrate into the Peru Coastal Current region, where the endemic S. bierri is to be found extending westwards to the west boundary of the Peru Coastal Current, and that both mesopelagic species, previously reported for the epipelagic level of this area, S. decipiens and Eukrohnia hamata, inhabit the upper mesopelagic level in the oceanic region, but also rise to the epipelagic level near the coast, where upwellings do occur. The lower mesopelagic levels (500 to 1000 m) are occupied by E. fowleri and S. macrocephala, this latter species being reported for the first time in the South Eastern Pacific Ocean. The other mesopelagic species, S. zetesios and S. bathypelagica, also found for the first time in this region, were not found in large enough numbers to obtain a correct view of their stratification. Eukrohnia bathyantarctica, described for the Southern Ocean and previously reported only in the bathypelagic levels of the Gulf of Mexico and Caribbean Sea, was found at only bathypelagic levels. The faunistic data from the R.V. A. Bruun Cruise 13, were compared with the corresponding hydrographical profiles.     

Factors affecting the flux of arsenic through the mussel Mytilus galloprovincialis

Radiotracer experiments were designed to study the effects of certain environmental and biological factors on arsenic accumulation and elimination processes in the Mediterranean mussel Mytilus galloprovincialis. Arsenic (as arsenate) uptake increased with increasing arsenic concentration in the water; however, the response was not proportional, indicating that accumulation was partially suppressed at higher external arsenic concentrations. In general, approximately 80% of the ⁷⁴As taken up was associated with the soft parts, with small mussels concentrating ⁷⁴As to a greater degree than larger individuals. The highest ⁷⁴As concentrations were recorded in the byssus and the digestive gland. Increased temperature enhanced both arsenic uptake and loss. Mussels in sea water at 19 S accumulated approximately three times more ⁷⁴As than those held at 38 S. Arsenic loss was much less affected by salinity, with only a tendency for greater arsenic retention noted at lower salinities. Studies carried out in the laboratory and in situ revealed that arsenic turnover was significantly more rapid in actively growing individuals living under natural conditions. Arsenic-74 loss from the in situ group was essentially biphasic, with biological half-times of approximately 3 and 32 days for the fast and slow compartments, respectively. The active secretion of arsenic in the byssal threads contributed to the total elimination of the element from the mussels.     

Individual effects and interactions of salinity,temperature, and zinc on larval development of the grass shrimp Palaemonetes pugio. I. Survival and developmental duration through metamorphosis

Larvae of the estuarine grass shrimp Palaemonetes pugio (Holthuis) were reared from hatch through successful completion of metamorphosis in 80 combinations of salinity (3 to 31%), temperature (20° to 35°C), and zinc (0.00 to 1.00 ppm Zn⁺⁺). Response-surface methodology was employed to depict the individual effects and interactions of the three factors on survival and developmental duration through total larval development. Outside the optimal salinity-temperature conditions of 17 to 27 S and 20° to 27°C, viability of larvae was reduced by both the individual effects of salinity and temperature and interactions between the two factors. Survival capacity of larvae and resistance adaptations to salinity and temperature were progresively reduced by zinc concentrations from 0.25 to 1.00 ppm Zn⁺⁺. Response-surface analysis of the data suggested that the duration of total larval development of P. pugio was least at salinities from 18 to 23 and at temperatures from 30° to 32°C. At both higher and lower salinity-temperature conditions and in increasing zinc concentrations from 0.25 to 1.00 ppm Zn⁺⁺, developmental rates were retarded. A significant zinc-temperature interaction existed, whereby increasing zinc concentrations reduced both survival and developmental rates of larvae more at suboptimal temperatures. Larval resistance to zinc toxicity was least at supraoptimal salinities, indicative of a significant zinc-salinity interaction. The reduced viability, restricted euryplasticity, and retarded developmental rates of P. pugio larvae developing in media with low-level zinc contamination would limit the distributive properties of the pelagic phase in the life cycle of the species and reduce recruitment both into and out of the parent estuarine population.     

Chemical aspects of mass spawning in corals. I. Sperm-attractant molecules in the eggs of the scleractinian coral Montipora digitata

This paper provides the first evidence for sperm chemotaxis in the Scleractinia. Montipora digitata Dana, 1845 (Scleractinia: Coelenterata) is a hermaphroditic coral which reproduces bi-annually, releasing egg-sperm bundles during the mass spawning at Magnetic Island (19°10S; 146°52E) in late spring-early summer, and autumn each year. The buoyant egg-sperm bundles float to the surface where they break apart, releasing eggs and sperm into the ocean. Fertilisation occurs after 30 min. Unfertilized eggs were collected, washed free of sperm, and freeze-dried. The eggs were extracted with dichloromethane, fractionated by chromatography on silica gel, and the fractions assayed for their ability to attract M. digitata sperm. The active fraction was further fractionated by high-performance liquid chromatography, resulting in the isolation of three highly unsaturated fatty alcohols: (1) dodeca-2,4-diynol; (2) tetradec-13-ene-2,4-diynol; (3) (14Z)-heptadeca-14,16-diene-2,4-diynol. Of these three compounds, only Compound 1 attracted sperm of M. digitata. Synthetic Compound 1, produced from simple precursors by known reactions, possessed sperm-attracting activity comparable to the naturally derived attractant. Preliminary experiments suggest that the natural mixture of Compounds 1, 2 and 3 in the ratio 1:4:9 is more effective in attracting sperm from M. digitata than sperm from other Montipora species. Sperm attractants may act to reduce the incidence of hybridisation between different species of Montipora.     

Reproductive effort of colonial and solitary breeding tree sparrows Passer montanus L.

A total of 250 nestboxes were arranged in five plots in a suburban area of Budapest, Hungary (19°04E, 47°41N). In each plot, 25 were placed at 50 m intervals to simulate solitary breeding and 25 3–5 m apart to simulate colonial breeding. Length of nest building period, feeding frequency, nestling mortality, nestlings' diet, productivity and parental condition were compared for colonial and solitary breeding tree sparrows Passer montanus. Parents with long nest-building periods, including the majority of first-year females, produced fewer young than parents which built over short periods. Parents fed nestlings morefrequently and nestlings had lower mortality in second than first broods; whether or not a third brood was reared was determined by the costs invested in first and second broods. Females that laid a third clutch had reared fewer young in first and second broods and were heavier than females that reared many young in two broods. Colonial birds had higher feeding frequencies, more similar diets and suffered lower nestling mortality than solitary parents for first broods, but they fed less frequently, diets were less similar, and nestling mortality was higher in second and third broods. It is suggested that colonial breeders benefited from the social stimulation of simultaneous feeding in first broods, but the advantage of synchronicity in feeding declined in second broods and the sparser breeding spacing of solitary parents was more advantageous for feeding in second and third broods. Birds that changed nest spacing between broods had fed nestlings less frequently and had higher nestling mortality before changing than birds which retained their spacing. Parents which changed from colonies to solitary nests fed more frequently with lower nestling mortality in the next brood than parents which retained colonial nests for their second (and third) brood. Solitary parents did not show such a relationship. The rearing of three broods caused higher weight loss in colonial than solitary parents.Correspondence to: L. Sasvári