Opportunistic Breeding Research Articles

Breeding phenology of Bufo viridis Laurenti, 1768 in Sicily

Bufo viridis Laurenti, 1768 is a common species that inhabits a wide variety of habitats. The different climates characterising its broad range lead to a high degree of variability in its seasonal activity and reproductive cycle. This paper reports some observations carried out on the breeding phenology of this species over a two year period in Mediterranean temporary ponds in Sicily. The reproductive period of Sicilian green toads extends into the autumn months, making it longer than that of other Italian populations. This behaviour seems due to the impact of xeric environmental conditions on the seasonal activity of the studied populations. The present study confirms that B. viridis is an opportunistic breeder with a wide margin of variability in annual reproductive cycle patterns, as would be expected of an ecologically variable species. The duration of the reproductive season varied between populations in the same year and between different years for the same population.

LH responses to single doses of exogenous GnRH in the Cape mole rat (Georychus capensis): the pituitary potential for opportunistic breeding

AbstractThe Cape mole rat Georychus capensis is a solitary mole rat that inhabits the winter rainfall region of the Western Cape Province of South Africa. Circulating basal concentrations of luteinizing hormone (LH) were found to be significantly higher in the breeding season in both sexes. During both the breeding and non‐breeding season, administration of exogenous gonadotropin‐releasing hormone (GnRH) increased circulatory LH levels. The magnitude of the LH response to an overdose of exogenous GnRH both in and out of the breeding season in males and females was not significantly different. Typically, seasonally breeding species exhibit a down‐regulation of the pituitary and reproductive functions out of the breeding season. It appears that there is no down‐regulation of GnRH receptors at the level of the pituitary out of the breeding season, because the pituitary responds to an exogenous GnRH challenge equally both in and out of the breeding period. The Cape mole rat exhibits the potential for opportunistic breeding out of the breeding period, provided that environmental factors are favourable. This finding questions whether this mole rat is actually a seasonal breeder or whether reproduction is hindered by the ecological constraint of the lack of opportunities to burrow and find mates at certain times of the year.

The pituitary potential for opportunistic breeding in the Cape dune mole-rat, Bathyergus suillus

In this paper we investigated the effect of sex and season on baseline and post-GnRH challenge luteinising hormone (LH) levels in a solitary, seasonally breeding mole-rat. Circulating basal concentrations of luteinising hormone (LH) were found to differ significantly with season in both sexes. However, no significant difference was found in circulating basal LH concentration between the sexes either within or out of the breeding season. The magnitude of the LH response to an exogenous pharmacological overdose of GnRH both in and out of the breeding season in males and females respectively was not significant. This finding suggests that there is no down regulation of GnRH receptors on the pituitary during the non-breeding season. Cape dune mole-rats thus have the potential for opportunistic breeding outside of the typical breeding period. We argue that this represents an adaptation to limited and brief opportunities for mating in this xenophobic and aggressive species.

Tonic Activation of Brain GnRH Immunoreactivity despite Reduction of Peripheral Reproductive Parameters in Opportunistically Breeding Zebra Finches

Opportunistically breeding species offer the unique opportunity to understand mechanisms in reproductive physiology that allow for extreme flexibility in the regulation of reproduction. We studied a well-known opportunistic breeder, the zebra finch (Taeniopygia guttata) to test the hypothesis that the reproductive axis of opportunists is in a constant state of ‘near-readiness’. In wild zebra finches, reproduction is highly correlated with rainfall, and in the laboratory, water availability and humidity are the strongest cues to affect reproductive activation. We therefore subjected individuals to water restriction for eleven weeks followed by a two week period of ad libitum access to water. The control group had water freely available for the entire experiment. We measured the state of activation of the hypothalamo-pituitary gonad (HPG) axis at three levels: in the hypothalamus by measuring immunoreactive (ir) cGnRH-I and cGnRH-II; in the anterior pituitary gland by measuring plasma luteinizing hormone (LH); and in the gonads by measuring gonadal volume and function. We found that water restriction caused a reduction in circulating LH concentrations and that testis volume was more likely to decrease in water restricted than in control birds. Subsequent short-term return to ad libitum water availability caused LH to return to baseline in water restricted birds. These changes occurred without significant changes in ir-cGnRH-I, ir-cGnRH-II, or in testis function. These data suggest that in these opportunistic breeders, an inhibition of parts of the reproductive axis is not necessarily correlated with full inactivation of reproductive potential. GnRH-ir cells in the hypothalamus appear to remain active and able to respond to subsequent stimulation.

Timing of reproduction in a Darwin's finch: temporal opportunism under spatial constraints

In seasonally unpredictable habitats, organisms cannot anticipate and prepare for a regular yearly reproductive period. Therefore, they opportunistically breed at any time of year. Opportunistically breeding birds are thought to maintain an activated reproductive system all year around. However, always being prepared can be costly and we therefore examined whether an opportunistic breeding strategy is realized in birds that are confined to harsh, climatically unpredictable habitats from which they cannot escape. We studied temporal variations in reproductive activity in small ground finches (Geospiza fuliginosa) in the Galápagos archipelago, Ecuador, by determining gonad sizes and plasma luteinizing hormone (LH) concentrations. We compared populations from the arid coasts of St. Fé and St. Cruz islands with birds from the St. Cruz humid highlands. The population from the arid St. Fé island increased gonad sizes and LH concentrations at least four months earlier in a wet (El Niño) year compared to the following dry year, indicating high temporal flexibility and opportunistic breeding. However, birds had completely regressed gonads for about 10 months between breeding events during a period of drought. Long‐term changes in environmental factors such as ambient temperature, daylength and light intensity did not appear to influence reproductive activity. Instead, environmental signals more directly connected to the occurrence of rains such as humidity, barometric pressure, or rainfall itself may stimulate reproduction. Our data demonstrate that small ground finches regulate the onset of reproductive activity flexibly like other opportunistic breeders. However, they differ from those by their unusual long‐term quiescence of the reproductive system during dry periods, which might minimize costs. Our findings suggest that within the continuum from seasonal to opportunistic breeding strategies, organisms evolutionarily optimize reproductive strategies to match local ecological conditions.

Overlapping litters and reproductive performance in the domestic rabbit

Rabbits ( Oryctolagus cuniculus), like many small mammals, have a marked postpartum estrus, and during the breeding season, are often both pregnant and lactating. We examined reproductive performance and allocation of resources by rabbit mothers with overlapping litters under presumably optimal conditions of unrestricted food, standardized litter size, and young reproductive age. Even under these conditions, females raising overlapping litters incurred higher costs in terms of greatly increased food intake and pup mortality than females raising nonoverlapping litters. Despite such costs, females with overlapping litters raised a larger total number of young to weaning than females without the load of simultaneous lactation and pregnancy. By more than doubling food intake during lactation, females maintained a stable baseline body weight throughout the study, suggesting that this, in a species with low fat reserves, might have priority over the short-term raising of a maximum number of young. Contrary to the expectation that females with overlapping litters would allocate more resources to their senior young, no appreciable difference in number, growth, or survival between pups of senior and junior litters was found either at birth or weaning. While this might reflect a strategy in this opportunistic breeder of distributing reproductive chances similarly between senior and junior litters, it is now necessary to investigate females' breeding strategy when resources are limited.

Opportunistic and multiple breeding attempts in plants and vertebrates of semi-deserts with unpredictable rainfall events through the year

Quantity and timing of rainfall vary among years in semi-deserts on the inland edges of Mediterranean-climate regions on all continents. One way that plants and animals might cope with the lack of predictability in rainfall is to have many fixed-season breeding attempts over a long lifespan, some of which are likely to succeed. Another approach is to breed sporadically, whenever conditions favour reproduction. In this review we compare breeding phenology of semi-desert organisms with that in other more predictable ecosystems. We test the hypotheses that breeding is unsynchronised within (a) communities and (b) species. We also examine evidence that opportunistic and multiple within-year breeding attempts in plants and animals is characteristic of semi-deserts with episodic rainfall that may occur at any time of the year. We conclude that opportunistic and iteroparous breeding occurs in many unrelated taxa, but that there are life-history, metabolic or dietary constraints to opportunism.

The diet of the Citril Finch (Serinus citrinella) in the Pyrenees and the role of Pinus seeds as a key resource

The diet of the Citril Finch (Serinus citrinella) in its eastern range is well known, especially in spring; however, no data are available from its western populations. The aim of this paper is to provide extensive data on the diet of the species in the Pyrenees. Samples were obtained by visual inspection of the gullet of 3394 birds from 49 different localities over an area of 4000 km2. Analyses revealed a clear phenological pattern: (1) the Arvensis period, November-January, when the bird relies on farmland seeds, mainlyChenopodium sp. (2) the Arboreal period, February-May, when the birds focus on Pine seeds (Pinus nigra andP. uncinata); (3) the Meadow-Ruderal period, June-October, when the bird feeds on several grasses and herbs, includingTaraxacum officinalis (mainly in June) andCirsium sp. (mainly August-September). Hence the Citril finch normally specialises in a few key species, but during certain periods may opportunistically expand its dietary range. Pine seeds appear crucial, both in determining the breeding distribution of the species and in allowing opportunistic spring breeding.

Flexible reproduction in wild canaries is independent of photoperiod

In temperate zone species, timing of the breeding season is determined by the minimum photoperiod or “critical day length” and the development of photorefractoriness. The integration of environmental cues like food, temperature, rainfall, and behavioural interactions determines the breeding window. However, there are a number of examples for breeding activities outside this time window. Here we investigated the possible mechanisms for early seasonal breeding activities in a population of wild canaries ( Serinus canaria) that inhabits an isolated island within the Madeiran archipelago. In December 1999, breeding activities were observed six weeks before the usual onset of the breeding season on the island, which can be related to unusual heavy rainfall during the two previous months. Moreover, testosterone (T) levels of the birds were significantly higher as during the same time in previous years and showed no difference to the T levels found in the breeding season. Thus, birds do not rely on the cues given by the photoperiod alone, but react independently of day-length to favourable environmental conditions, like water availability. Therefore, the individuals of this wild canary population can be considered as seasonal opportunistic breeders.

Opportunistic breeding in the Cape spiny mouse (Acomys subspinosus)

The relationship between Protea humiflora and its small mammal pollinators was investigated at the fynbos/karoo ecotone in the Riviersonderend Mountains, South Africa. Acomys subspinosus occurred in low densities at Jonaskop, with around 3.9 ± 2.0 (mean ± 1 S.D.) individuals/ha (six months of trapping). Low repeat trapping may result from low survivorship or emigration from the area after flowering of P. humiflora. Mean mass of A. subspinosus was lower than published values due to a high proportion of juveniles in the population, coinciding with P. humiflora flowering. Comparison with other published data suggests that A. subspinosus is an opportunistic breeder. This contrasts with its geographically nearest congener, A. spinosissimus, in which breeding is restricted to the summer months, but is similar to data for other Acomys species. Minimum home range areas (≈0.23 ha, minimum overnight distance 39 ± 6 m), were relatively high and may indicate a species surviving on thinly dispersed resources.

Reproductive Endocrinology of a Free-Living, Opportunistically Breeding Passerine (White-Winged Crossbill, Loxia leucoptera)

The aim of the study was to investigate, in a free-living opportunistic breeder, the White-winged Crossbill (Loxia leucoptera), the degree to which annual changes in photoperiod and differences in the timing of breeding in different years correlate with the secretion of reproductive hormones. Seasonal changes in concentrations of luteinizing hormone (LH) and prolactin were compared in White-winged Crossbills in interior Alaska (64°50′N, 147°50′W) in plasma samples taken in May–December 1994 and July 1998–August 1999. The birds were in breeding condition in May–July 1994 and in February–April 1999. Additional observations were made on plasma testosterone (T) in males in 1998/1999. In 1998, in adult males, plasma LH and T were low in September–October (LH: <0.4 ng/ml; T: undetectable) and increased significantly in November (LH: 3.65 ± 2.20 ng/ml) and December (T: 0.69 ± 0.88 ng/ml). Thereafter, plasma T increased to 2.35 ± 2.03 ng/ml in March/April and returned to basal levels by June/July. In 1998/1999, in females, plasma LH did not change seasonally. In males in 1994, plasma LH was higher in June (2.15 ± 1.22 ng/ml) and July (2.86 ± 0.69 ng/ml) than in the same period in 1999 (June: 1.20 ± 1.82 ng/ml; July: 1.37 ± 1.03 ng/ml), which is consistent with the later breeding season in 1994. In 1994 and 1998/1999 in both sexes, concentrations of plasma prolactin were elevated when day lengths exceeded about 12 h, being high during late spring to early fall (males: 40–130 ng/ml; females: 40–140 ng/ml) and low during winter (<15 ng/ml in both sexes). In 1994 and 1998, molt occurred during the fall, at a time when breeding did not occur, although conifer seeds were sufficiently abundant to support a large bird population. These data are consistent with the view that White-winged Crossbills have the potential to breed at all times of the year except when molting in the fall. It is likely that a photoinduced increase in prolactin secretion is a factor that induces molt. Crossbills may not breed when they are molting because of increased metabolic demands for feather formation and for the maintenance of basal metabolism at a time of year when ambient temperatures fall well below 0°. Alternatively or in addition, the birds may become reproductively photorefractory.

Photoperiodic control of seasonality in birds.

This review examines how birds use the annual cycle in photoperiod to ensure that seasonal events--breeding, molt, and song production--happen at the appropriate time of year. Differences in breeding strategies between birds and mammals reflect basic differences in biology. Avian breeding seasons tend to be of shorter duration and more asymmetric with respect to changes in photoperiod. Breeding seasons can occur at the same time each year (predictable) or at different times (opportunistic), depending on the food resource. In all cases, there is evidence for involvement of photoperiodic control, nonphotoperiodic control, and endogenous circannual rhythmicity. In predictable breeders (most nontropical species), photoperiod is the predominant proximate factor. Increasing photoperiods of spring stimulate secretion of gonadotropin-releasing hormone (GnRH) and consequent gonadal maturation. However, breeding ends before the return of short photoperiods. This is the consequence of a second effect of long photoperiods--the induction of photorefractoriness. This dual role of long photoperiods is required to impart the asymmetry in breeding seasons. Typically, gonadal regression through photorefractoriness is associated with a massive decrease in hypothalamic GnRH, essentially a reversal to a pre-pubertal condition. Although breeding seasons are primarily determined by photoperiodic control of GnRH neurons, prolactin may be important in determining the exact timing of gonadal regression. In tropical and opportunistic breeders, endogenous circannual rhythmicity may be more important. In such species, the reproductive system remains in a state of "readiness to breed" for a large part of the year, with nonphotic cues acting as proximate cues to time breeding. Circannual rhythmicity may result from a temporal sequence of different physiological states rather than a molecular or cellular mechanism as in circadian rhythmicity. Avian homologues of mammalian clock genes Per2, Per3, Clock, bmal1, and MOP4 have been cloned. At the molecular level, avian circadian clocks appear to function in a similar manner to those of mammals. Photoperiodic time measurement involves interaction between a circadian rhythm of photoinducibility and, unlike mammals, deep brain photoreceptors. The exact location of these remains unclear. Although the eyes and pineal generate a daily cycle in melatonin, this photoperiodic signal is not used to time seasonal breeding. Instead, photoperiodic responses appear to involve direct interaction between photoreceptors and GnRH neurons. Thyroid hormones are required in some way for this system to function. In addition to gonadal function, song production is also affected by photoperiod. Several of the nuclei involved in the song system show seasonal changes in volume, greater in spring than in the fall. The increase in volume is, in part, due to an increase in cell number as a result of neurogenesis. There is no seasonal change in the birth of neurons but rather in their survival. Testosterone and melatonin appear to work antagonistically in regulating volume.

Effects of humidity on reproductive behavior in male and female zebra finches (Taeniopygia guttata).

Zebra finches (Taeniopygia guttata) are regarded as opportunistic breeders with reproductive behaviors mediated by short-term proximal environmental conditions. This article provides experimental evidence for the role of humidity in reproductive behaviors. Zebra finches were subjected to experimentally manipulated high levels of relative humidity. Males gathered more nest material and sang more. Females showed no relationship between humidity and gathering nest materials, egg laying, or changes in rates of vocalization.

Seasonal Changes in Brain GnRH Immunoreactivity and Song-Control Nuclei Volumes in an Opportunistically Breeding Songbird

White-winged crossbills (Loxia leucoptera) are opportunistic breeders that can nest at almost any time of year if there is sufficient food. Other cardueline finches that have been shown to breed on a strictly seasonal schedule become absolutely refractory to the stimulatory effects of long-day photoperiod, dramatically down-regulate hypothalamic gonadotropin-releasing hormone (GnRH), and reduce the volume of several song-control nuclei in autumn. This study examined whether changes in photoperiod modify the GnRH and song-control systems in white-winged crossbills. Adult male and female crossbills were captured and held on a naturally changing photoperiod. Brains of male and female birds were collected in May, October, and January. GnRH content was assessed by immunocytochemistry and the volumes of Nissl-defined song-control nuclei (HVc, Area X, and the robust nucleus of the archistriatum) were reconstructed. In contrast to other cardueline finches, GnRH immunoreactivity was relatively stable across the year, exhibiting only modest seasonal variation. The song control system, on the other hand, exhibited large seasonal changes as well as sex differences. Thus, crossbills appear to maintain hypothalamic GnRH content year round, perhaps to facilitate a rapid response to favorable breeding conditions, even on short days. However, song control nuclei are dramatically affected by photoperiod. Future work should examine these systems in crossbills breeding on short days to compare photoperiod-dependent and -independent effects on neural plasticity.

Photoperiodic Regulation of the Reproductive Axis in Male Zebra Finches, Taeniopygia guttata

In arid central Australia, breeding of zebra finches (Taeniopygia guttata) tends to take place immediately after the aperiodic rainfall—this being irrespective of the time of year. As a result, zebra finches have traditionally been considered nonphotoperiodic. Despite this, there are some published reports of photoperiodic effects on behavior in zebra finches. Therefore, we transferred singly housed male zebra finches from a photoperiod of 14 h light and 10 h dark per day (14L:10D) to either 20L:4D or 8L:16D. Control birds remained exposed to 14L:10D. Plasma LH, testicular volume, and body mass were assessed at the start of the experiment and at intervals for a period of 56 days. Testicular mass was measured at the end of the 56-day period. Plasma LH increased significantly in the 20L:4D group after 14 days, but decreased again by 56 days, presumably an effect of increased gonadal steroid negative feedback. Plasma LH did not change significantly in the other two groups. Testicular volume increased steadily in the 20L:4D group during the treatment period and it was significantly higher than that of the 8L:16D group at the end of the experiment. After 56 days of treatment, combined testicular mass in the 20L:4D group was much greater than that of the 8L:16D group, but not quite statistically different from the 14L:10D group. There was no statistical difference in testicular mass between the 14L and 8L groups. Body mass did not differ between any of the groups at any time in the experiment. Our results are consistent with zebra finches being photoperiodic to some degree, despite their opportunistic breeding strategy. When considered in conjunction with recent reports of photoperiodic responses in tropical avian species, these data suggest that the ability to respond to changing photoperiod is more common among avian species than previously hypothesized.

Torpor patterns in the pouched mouse (Saccostomus campestris; Rodentia): a model animal for unpredictable environments.

Patterns of spontaneous and induced daily torpor were measured in the Afrotropical pouched mouse (77-115 g), Saccostomus campestris, in response to photoperiod, temperature, and food deprivation, using temperature telemetry. Photoperiod had no influence on the incidence, depth, or duration of daily torpor in either males and females. Although the testis size index decreased in response to food deprivation and photoperiod by a maximum of 24%, full testis regression did not occur. Torpor bout duration was, on average, 5.3 h, independent of photoperiod and ambient temperature. Males did not enter torpor in response to food deprivation but did in response to low ambient temperature, though significantly less frequently than females. At normothermia, the body temperatures (daily minimum, mean, maximum) of males were significantly lower than those of females. Minimum body temperatures of both males and females during torpor did not fall below 20 degrees C at an ambient temperature of 15 degrees C. The patterns of torpor measured here differ from those observed in species from strongly seasonal environments. They suggest adaptation to an environment rendered unpredictable by the El Niño Southern Oscillations. As an aseasonal, opportunistic breeder capable of year-round adaptive hypothermia, the pouched mouse represents an excellent model animal for research on physiological and behavioral adaptations to unpredictable environments.

Population Dynamics and Movement Patterns of Bolam&amp;apos;s Mouse, Pseudomys bolami, at Roxby Downs, South Australia.

Basic ecological data were collected on Bolam&amp;apos;s mouse, Pseudomys bolami, during a six year trapping study at Roxby Downs in northern South Australia. Pseudomys bolami inhabited mixed arid land systems in the Roxby Downs area, sheltering in the longitudinal orange sand dunes but foraging at night principally on the adjacent chenopod swales. Distances of up to 334m were traversed in a single night and recapture rates were high within trapping sessions but low between sessions suggesting wide-ranging movements and short-term residency. Dietary observations imply an omnivorous diet. Whilst the closely related P. hermannsburgensis is regarded as having an opportunistic breeding strategy, P. bolami reproduced opportunistically but also regularly in spring and early summer. Different reproductive strategies may be influenced by the latitudinal distribution of P. bolami near the southern margin of the arid-zone, where rainfall can occur at any time of year but winter rainfall is more predictable. This contrasts with P. hermannsburgensis, which inhabits more northerly arid areas where rainfall is erratic and unpredictable. There was a 10 fold fluctuation in P. bolami captures over the trapping period with numbers of P. bolami and house mice, Mus domesticus peaking 12 months after above average rains during 1992. Rodent captures remained high for two years afterwards. Although P. bolami persisted throughout the study period, M. domesticus was only present when conditions were above average. Trap success of P. bolami or M. domesticus did not vary between the five sub habitats at the 1 ha study site. However, trap success of P. bolami was positively related to vegetation cover, particularly cover of low bluebush, Maireana astrotricha. Maireana astrotricha may be important in providing both food and cover for P. bolami, a scenario consistent with the utilisation of spinifex, Triodia spp. by P. hermannsburgensis in the northern arid-zone. Past over- grazing of chenopod shrublands may have reduced cover and contributed to the suggested decline in the distribution of P. bolami.

The importance of green seed in the nitrogen nutrition of the Zebra Finch Taeniopygia guttata

Abstract Australian grass‐finches are widely reported to consume large quantities of green seed when it becomes available, and the opportunistic breeding of wild Zebra Finches in the arid zone has been correlated with the occurrence of rain. In this study, green and ripe seeds were harvested from seven pasture and weed grasses grown in experimental plots and, along with three cereal flours and whole‐egg powder, were analysed for the amino‐acid composition of their protein. The relative levels of ten amino acids essential in the diets of growing birds were compared between samples using a cluster analysis dendrogram generated from Raabe's Similarity Index. The protein of all green seeds clustered with whole egg, and away from all but one of the ripe seeds and seed products. Green and ripe seed profiles were found to be significantly different by a two‐sample multivariate test of significance (Hotelling's T2). Histidine, lysine, phenylalanine and threonine were the amino acids most different. Of these four amino acids, lysine and threonine (along with methionine) were potentially limiting in ripe seeds when compared with whole‐egg protein. In green seeds, lysine was only marginally limiting, threonine was no longer limiting, but methionine was still limiting when compared with whole‐egg protein. These results indicate that the benefit of green vs ripe seed in the diet of breeding Zebra Finches is partly a higher level of the limiting essential amino acid, lysine, and partly a higher intake and throughput of soft green seed and consequent greater extraction of limiting essential amino acids.

New findings in body mass regulation in zebra finches (Taeniopygia guttata) in response to photoperiod and temperature

Small birds in temperate zones increase body mass in winter (Lehikoinen, 1987). We investigated daily and annual variation of body mass and fat reserves of locally‐reared Australian zebra finches Taeniopygia guttata, by exposing them to different photoperiods, feeding periods, and temperatures.Experiments with long and short photoperiods and long and short feeding periods, showed that long photoperiods increased body mass and fat reserves of the zebra finches, and readiness to breed, independently of the actual feeding period.Furthermore, the zebra finches in indoor aviaries with constant temperature (22‐24°C) and in outdoor aviaries with ambient temperature, both exposed to the natural daylength changes of Bielefeld, Germany (52 °N), had high dawn body mass in summer (12.9 and 12.0‐12.4g, respectively) and low in winter (10.7 and 11.1 g, respectively). Thirty to sixty percent of these mass changes were related to changes in fat reserves, so that the finches had only 0.1‐0.2g of metabolizable fat reserves in short photoperiods (or in winter), which increased up to 1.5g in long photoperiods (or in summer).Indoor finches consumed more seeds in summer than in winter (3.3 vs. 2.7g/day), while outdoor finches consumed 4‐5g of seeds per day throughout the year, which probably represented the limit of energy intake for a 11‐13g bird (Kirkwood, 1983). Nightly mass loss, increasing from 0.7g in summer up to 2.0g in winter, was highly positively correlated with night length, not influenced by ambient temperature. Foraging before dawn and after dusk, roosting with well‐filled crops, and decreasing body mass and fat reserves, seem to be adaptations of zebra finches for survival in winter. The summer fattening probably accelerates reproduction in this opportunistic breeder, by allocating more time to reproductive behaviour and more endogenous nutrients to egg‐formation.

The Role of Androgens in Opportunistic Breeding, Tropical Frogs

In this paper we present the results of an interspecific study on the androgen levels of several species of Old World tropical frogs. These data are used to test three hypotheses: (1) in unpredictable, aseasonal environments with opportunistic breeding, gonadal hormones have a permissive rather than an activating role in the expression of mating behavior (Crews and Moore, 1986), (2) a behaviorally induced androgen response is present in opportunistic aseasonal species as well as in temperate seasonal breeders (Crews and Moore, 1986; Wingfieldet al.,1990), and (3) because maintenance of high plasma testosterone levels has an associated energetic and/or fitness cost (Moore and Marler, 1987; Wingfieldet al.,1990), continuously breeding frogs have lower testosterone levels than seasonal, discontinuous breeders. Androgen and corticosterone levels were found to vary significantly among species exhibiting the same mating behavior, providing support for the hypothesis that hormones have a permissive role. Differences in androgen level were correlated with (1) variation in level of male–male aggression, (2) relative density of male frogs, and (3) call rate and power output of male vocalization. These results indicate that a behaviorally induced androgen response is present in opportunistic, aseasonal breeders. Furthermore, tropical, aseasonal, opportunistic breeding frogs appeared to have lower androgen levels, generally, than confamilial temperate, seasonal breeders. This difference is consistent with the hypothesis that maintenance of high plasma testosterone levels is energetically expensive.