Effects of inter-pond distance on the breeding ecology of tungara frogs

Habitat and resource distributions can influence the movement and aggregation of individuals and thus have important effects on breeding behavior and ecology. Though amphibians have been model systems for the study of breeding behavior and sexual selection, most studies have examined breeding behavior within a single pond. As a result, little is known about how inter-pond distance affects breeding amphibians. We studied the effects of inter-pond distance on the breeding ecology of the tungara frog, Physalaemus pustulosus, in replicated pond arrays in which distance was varied from 0 to 50 m. We predicted that male site fidelity and male aggregation within arrays would increase with inter-pond distance, and that the opportunity for mate choice and oviposition site selectivity by females would decrease with the distance between ponds. Male site fidelity did increase with inter-pond distance. However, male aggregation decreased with distance, such that males tended to be more evenly spaced among ponds when ponds were farther apart. The opportunity for mate choice by females, measured as the number of males within the phonotactic radius of females, also decreased with inter-pond distance. Each of these three responses was consistent with a threshold effect between 5 m and 10 m in inter-pond distance. This threshold corresponded to the maximum distance at which females in laboratory choice experiments exhibited phonotaxis toward the ”whine” call of a tungara male, suggesting that phonotactic limits may play an important role in tungara movements and spacing patterns. The distribution of egg masses among ponds, a potential correlate of oviposition site selectivity, did not vary with inter-pond distance. Multiple egg masses deposited on the same night were significantly overdispersed in all distance treatments, implying that females may select oviposition sites to avoid conspecific egg masses over distances of at least 50 m. Collectively, these results demonstrate that inter-pond distance may indeed affect amphibian breeding and movement behavior, and that consideration of multiple habitat patches and their spatial distributions can provide new insights into even the most well-understood mating systems. Received: 9 January 1999 / Accepted: 1 August 1999     

Male breeding behaviour and mate acquisition in the Common toad, Bufo bufo

The breeding behaviour of a Common toad ( Bufo bufo ) population was studied at a breeding pond on Portland, Dorset. Major toad movements towards the breeding pond occurred on warm, wet nights when the temperature did not fall below 6°C. Males arrived earlier than females resulting in an initially exaggerated sex ratio which then slowly reduced as the breeding season progressed. No size assortative pairing was found although large males were more successful at mating than small males. Although males demonstrated no ESS (evolutionary stable strategy) in searching for a female between sites within the pond evidence suggesting the existence of one between the pond and an undefined area surrounding it was found.     

大凉螈繁殖生态

大凉螈是我国特有的珍稀有尾两栖动物,其种群数量目前呈现明显下降趋势,然而涉及该物种保护的繁殖生态学研究仍十分匮乏。通过融合围栏陷阱及标志重捕的样方调查法,对大凉螈石棉栗子坪种群繁殖个体和变态登陆幼体的迁徙、繁殖群体种群大小、繁殖场内雌雄有效性比变化等进行了研究。运用Jolly-Seber法估测了繁殖种群大小,运用单因素方差分析或Kruskal-Wallis秩和检验比较了不同时期进入繁殖场的雄性大凉螈头体长及体重,运用t检验或者Wilcoxon秩和检验比较了雌雄性间形态上的差异,运用t检验、t′检验或Wilcoxon秩和检验比较了野外抱对雄性与非抱对雄性间的体征差别,运用Pearson相关分析探讨了雌性产卵量与其身体形态的关系,同时观察了卵的孵化情况。研究结果表明:大凉螈的繁殖季为每年的4月下旬到7月下旬,幼体最早于8月上旬变态登陆。估测调查地繁殖场内雄性大凉螈繁殖种群大小约为391尾,雄螈较雌螈更早进入繁殖场且在场内停留时间更长,体重较轻的雄螈较晚迁入繁殖场。有效性比明显偏雄(雌/雄:0.03—0.10)。雌雄间具明显性二型性,雌性个体的头体长、体重及肥满度均大于雄性,而雄性的尾高和尾长占全长的比例则大于雌性。对比自然抱对雄性和非抱对雄性个体发现,抱对个体在头体长、体重和尾高等体征方面显著大于非抱对个体,暗示这些形态特征可能在雄性竞争配偶的过程中起到关键作用。雌螈在室内条件下平均产卵数为176枚,产卵历时2—4 d,产卵量与雌性肥满度正相关,卵的平均孵化期为15.7 d,孵出幼体平均全长为9.74 mm。     

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

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

Patterns of roost site selection and use by Southern Ground-Hornbills in north-eastern South Africa

Different habitats may be used for the needs of various aspects of an animal’s life. Southern Ground-Hornbill Bucorvus leadbeateri groups announce their presence within year-round territories by calling at dawn from their overnight roost sites. Knowledge on ground-hornbill roosting habits is limited. Groups roost in large trees, apparently close to where they end up after daily foraging. We investigated patterns of roost site selection and use for four Southern Ground-Hornbill groups in the Associated Private Nature Reserves, north-eastern South Africa, based on data from GPS-satellite transmitters. The number of roost sites used per month averaged 15.4 ± 4.7 across all groups, indicating little evidence of strong preferences for specific sites. This number was least when groups were breeding, decreasing throughout the early wet season (October–December) and was lowest during the late wet season (January–March) when actively breeding groups frequently roosted close to the nest (54–83% of roosts <1 000 m of the nest). As might be expected, the mean monthly number of nights per roost peaked during the breeding season (December–January). Riparian habitats were preferred for roosting during the breeding season, whereas disturbed areas, as well as Combretumand mopane-dominated habitats were preferred during the dry non-breeding season. Adequate large trees not only for nesting, but also for roosting, particularly in riparian habitats, may therefore be an important and potentially limiting factor for the successful reproduction of Southern Ground-Hornbills.     

Social regulation of plasma estradiol concentration in a female anuran

The behavior of an individual within a social aggregation profoundly influences behavior and physiology of other animals within the aggregation in such a way that these social interactions can enhance reproductive success, survival and fitness. This phenomenon is particularly important during the breeding season when males and female must synchronize their reproductive efforts. We examined whether exposure to conspecific social cues can elevate sex steroid levels, specifically estradiol and androgens, in female túngara frogs (Physalaemus pustulosus). We compared plasma estradiol and androgen concentrations in wild-caught females before and after exposure to either natural mate choruses or random tones. After exposure to mate choruses for 10 consecutive nights, estradiol concentrations were significantly elevated whereas there was no significant elevation in estradiol concentrations in the group of females exposed to random tones for 10 nights. Plasma androgen concentrations were not significantly changed after exposure to either natural mate choruses or random tones for 10 consecutive nights. Social modulation of estradiol concentrations may be important in maintaining a female's reproductive state while males are chorusing. To our knowledge, this is the first study to demonstrate social regulation of estradiol concentration in female anurans.     

Life stage specific variation in the occupancy of ponds by Litoria aurea,a threatened amphibian

Breeding aggregations are a reproductive strategy to increase mate finding opportunity. However, because aggregations skew the distribution of mature animals through conspecific attraction, rather than resource availability, the distribution of breeding sites may be reduced, so that not all suitable breeding sites are used. To examine the relationship between landscape and reproductive strategies of a threatened frog, Litoria aurea, we studied its distribution at Sydney Olympic Park over two breeding seasons. We aimed to: (i) determine the distribution and predictors of breeding ponds; and (ii) assess the significance of dispersal in the juvenile age‐class. We found that the distribution of the calling males was highly skewed and occurred in large, well‐connected ponds. Despite this, breeding ponds were not aggregated; pond size was the single factor explaining the distribution of breeding ponds. Juvenile frogs dispersed from breeding ponds and were not associated with a specific pond characteristic. Less breeding occurred in the second season during which fewer ponds were used for breeding including many different ponds from the previous year. These changes suggest that breeding effort and breeding pond choice are dynamic and therefore knowledge of the factors that drive breeding events will be a powerful tool in managing species, particularly in light of changing climatic regimes.     

Mating System Variation in Woodhouse's Toad (Bufo woodhousii)

Sexual selection and mating-system variation was studied in two disjunct populations of Woodhouse's toad (Bufo woodhousii) with dissimilar breeding-period durations. In spite of the differences in breeding-period duration, nightly operational sex ratios were roughly equivalent and strongly male biased in both populations, ranging from 0 to 0.5 in the population with a prolonged breeding period, and from 0 to 0.2 in the explosively breeding form. Although operational sex ratios were similar, some males mated with up to three females in the population with an extended breeding period, whereas none of the males mated polygynously in the population with a short breeding season. Two alternative reproductive tactics, satellite and active-searching behavior, were only exhibited by males under high densities in the population with a short breeding period. In the population with an extended breeding period, male mating success was positively related to call rate, but independent of size or number of nights of participation in chorus activity. Although operational sex ratios were equally low, mating was random by size, number of nights of participation in chorus activity, and call rate for males in the explosively breeding population.     

Number of males in primate groups: Comparative tests of competing hypotheses

Primate social groups frequently contain multiple males. Male group size has been hypothesized to result from male mating competition, but the selective factors responsible for the evolution of multimale groups are unclear. Short breeding seasons create situations that are not conducive for single males to monopolize mating access to females, and may therefore favor the formation of large male groups. Alternatively, since the costs of mate defense increase with the spatial clumping of females, female group size may be a primary determinant of the number of males in a primate group. We used comparative methods designed to control for the potentially confounding effects of hidden third variables associated with phylogeny to test the breeding season and female group size hypotheses for the evolution of multimale groups. Our results revealed no association between breeding season duration and the number of males in groups. In contrast, we provide support for the female group size hypothesis by demonstrating a strong pattern of correlated evolution between female and male group size. © 1996 Wiley-Liss, Inc.     

Non-random mating by size in American toads, Bufo americanus

Male American toads, Bufo americanus, captured in amplexus at least once during the season were significantly longer than males captured only singly in the same breeding population in all three cases studied. Absence of any correlation between the body lengths of the members of amplexing pairs argues against selection for optimal relative size. Males successful at breeding were those that (1) were larger, (2) were at the pond more nights, and (3) spent more time calling than unsuccessful males. Residency, per se, did not enhance breeding success, however, as on any given day successful males were equally likely to come from recaptured males previously present at the pond and unmarked males presumed new to the pond. Amount of movement about the pond did not vary with breeding success; all males tended to remain in one area on any given night and to change areas between nights. A very low displacement rate (about 7%) of males in amplexus argues against primacy of this form of male-male competition in producing differential mating by size. Rather it appears that some combination of other forms of intrasexual competition and female choice caused the mating pattern seen.     

Plasticity in the mating system of the longnose filefish,Oxymonacanthus longirostris, in relation to mate availability

The mating system of the longnose filefish,Oxymonacanthus longirostris, was examined on coral reefs of Okinawa, Japan. This species has been shown previously to be monogamous. Fish were usually found swimming together in heterosexual pairs with the male and female sharing the same feeding territory. However, both monogamous and polygynous (bigamous) males were found in the present study. Polygynous males, which were larger than monogamous males, visited and stayed several minutes in turn with each female within the territories. Although most males were monogamous in the early breeding season, over 20% of males mated polygynously in the late breeding season. The adult sex ratio in the former was unbiased, but became slightly female-biased toward the end of the breeding season because of the higher disappearance rate of males. The higher disappearance rate may be due mainly to a higher mortality rate of males resulting from a greater deterioration of physical condition during the breeding season. Thus, the mating system varied with the change of the adult sex ratio. Plasticity in the mating system of this species may be the outcome of male mating tactic depending on local mate availability.     

Immunological aspects of gestation in the tammar wallaby, Macropus eugenii.

Sensitization to male histocompatibility antigens and repeated pregnancy to the same male were found to have little effect on fertility or length of gestation in the tammar wallaby, M. eugenii. However, in some sensitized females a long interval occurred between removal of pouch young and the next birth. In addition to studies on fertility, the immunological response of female tammars to their mate has been examined by one-way mixed leucocyte culture (MLC) carried out at the beginning and end of one breeding season. In virgin females, examined at the beginning of the breeding season, the MLC response to the prospective mate peaked on day 6. In contrast, by the end of the season, MLC responses were much lower and peaked earlier, on day 3 to day 5.     

Spatial and Temporal Distribution of Song in the Yellowhammer Emberiza citrinella

The temporal and spatial distribution of song was studied in a population of yellowhammers Emberiza citrinella. Song was most frequent during the breeding season, and within the breeding season during the fertile period of both first, second, and replacement clutches. Song activity peaked at sunrise and sunset. During the fertile period most singing took place in the central parts of the territory. Song post heights peaked during the fertile period, and more song posts lacked foliage at that time. Intrusions by male conspecifics peaked in the fertile period and in territories where males sang relatively little. Song activity and mate guarding were strongly positively correlated. Song volume was loud and song was thus apparently used in long-distance communication. These observations are in accordance with a male deterrence hypothesis, suggesting that males sing to deter neighbouring males from trespassing during the fertile period of their mate. A female attraction hypothesis, suggesting that males sing to attract neighbouring females and thereby obtain extra-pair copulations, and a female reproduction hypothesis, suggesting that males sing to start the female reproductive cycle, were partly supported by observations.     

The Behavioral Ecology of Libellula luctuosa (Burmeister) (Anisoptera: Libellulidae): I. Temporal Changes in the Population Density and the Effects on Male Territorial Behavior

A single population of a common pond dragonfly, Libellula luctuosa, was studied at a site where the density of males increased dramatically during the breeding season. Early in the summer one active male was found on each territory on the pond. Satellite males were only occasionally found on the territories. Later in the season the number of males per territory increased so that two or more males simultaneously defended on many of the territories, and several satellite males occupied each of the territories. The number and rate of female visitations per day did not change over the summer. These factors resulted in a change in the operational sex ratio with variations in male density. Male behavior was also altered with increasing population density. As male density increased, males were less likely to be seen perching on their territories and more likely to be seen performing aggressive acts such as chasing nearby territorial males and chasing intruders. At high male density, the duration of territorial behaviors was shorter than at low male density. Thus, the percent of a time budget spent in any one activity did not change despite the change in number of males present. Male activity in L. luctuosa is not strictly determined by the opportunity for aggression. Costs of aggression associated with territoriality are minimized by maintaining flexible territorial behaviors.     

Variation in Song Rate during the Breeding Cycle of the Chaffinch,Fringilla coelebs

The variation in song rate during the breeding season was studied in two individually marked chaffinch Fringilla coelebs populations. We gathered data to investigate especially the recently presented mate-guarding hypothesis. The active singing has been supposed to function as a form of mate guarding during the female's fertile period by announcing the high status of the male and preventing extra-pair copulations by neighbouring males. There was no clear dawn chorus in the chaffinch, i.e. a peak in the song rate before sunrise. Male chaffinches continued to sing after mating, but the song rate dropped significantly. In contrast to the mate-guarding hypothesis the song rate was lower during the fertile period of the female than during pre-mating and incubation. Thus, the males do not announce the fertility status of their mates or their own quality and status by active singing. The song does not function as a form of mate guarding in the chaffinch. One function of the song of the chaffinch is mate attraction: singing activity was highest before pair formation in early spring and decreased after mating but increased again if the male lost his mate later in the breeding season.     

The effect of mate change and new partner's age on reproductive success in the barnacle goose, Branta leucopsis

Mate retention frequencies and correlations between mate changeand reproductive performance were estimated in a populationof barnacle geese, Branta leucopsis, breeding on the islandof Gotland in the Baltic. About 90% of the pairs remained togetherfrom one breeding season to the next. Only 2.4% of the pairsdivorced, most mate changes being consequences of the deathof one partner. Divorces were not forecast by low reproductivesuccess, and seemed to be accidental. In the season before matechange, there was no difference in reproductive performance,measured as clutch size, hatching date, and number of fledgedyoung, between faithful pairs and pairs where one partner wassubsequently changed. However, in the first season with a newpartner, clutch size and number of fledged young decreased onaverage. Hence, because mate change led to a reduction in reproductivesuccess, it was concluded that mate retention is advantageous.Our results suggest that this reduction is more likely due tothe lower average age or breeding experience of new partnersthan to the benefits of breeding experience with one particularpartner. [Behav Ecol 1991 ;2:116–122]     

Breeding Tubercles, Papillomatosis and Dominance Behaviour of Male Roach (Rutilus rutilus) During the Spawning Period

We studied the relationship between the breeding tubercle ornamentation (i.e. skin roughness) and male pre‐spawning dominance and courtship behaviour in roach (Rutilus rutilus) within an experimental laboratory system. Sexually mature fish were caught during their migration to their spawning pond and their behaviours were studied in an artificial spawning arena. Males behaved naturally both in terms of male–male interactions and attempts to achieve spawnings. Males having many, large breeding tubercles (i.e. rough skin) were significantly more often dominant in our dyadic trials than those with smooth skin. The dominant male in the trial exhibited a more active courtship behaviour than its subordinate rival. Papilloma skin disease did not affect the dominance rank. As a result of the relationship between skin roughness and male dominance, breeding tubercles may be used by the females as a cue for choosing a high‐quality mate in a roach lek. Thus, breeding tubercles might offer a workable tool for examination of sexual selection among cyprinids.     

Seasonal Variation in Mate Choice of Photinus ignitus Fireflies

Mate choice by either sex may vary with changes in the associated costs and benefits, determined by factors such as the availability of potential mates and variation in mate quality. We examined seasonal variation in operational sex ratio, courtship behavior, spermatophore mass, egg count, and the relationship between morphological traits and mating success in Photinus ignitus fireflies to determine if mate choice in either sex varied with the availability and relative reproductive investment of fertilizable females and sexually active males. Successfully mating males had larger lanterns than unsuccessful males when the operational sex ratio was male‐biased. In addition, female responsiveness to male signals increased as the number of courting males decreased, and male spermatophore mass decreased with body size across the mating season. Successfully mating females had larger body mass than unsuccessful females. Female body mass predicted egg count and female rejection by males increased as the season progressed and female size decreased. These results suggest that both male and female P. ignitus exhibit mate choice, and that such choice is influenced by seasonal variation in the abundance and quality of potential mates.     

Testis size depends on social status and the presence of male helpers in the cooperatively breeding cichlid Julidochromis ornatus

Cooperatively breeding animals, in which helpers may participatein reproduction with dominant breeders, are ideal species forexamining intraspecific variation in testis size because theyoften exhibit both monogamous breeding (low risk of sperm competition)and polyandrous breeding (high risk) within a population. However,little is known about testis investment as a result of spermcompetition in these animals. The substrate-brooding cichlidfish Julidochromis ornatus has a cooperatively breeding system,in which some males mate monogamously and other males reproduceas dominant breeders or helpers within cooperatively breedinggroups, in which male helpers frequently sire young. We examinedthe relationship between testis investment and male social statusin relation to the risk of sperm competition. As predicted fromsperm competition models, in groups with male helpers, boththe male breeders and the male helpers invested more in testesmass, compared to breeding males without male helpers. We alsofound a positive relationship between the testes mass of malebreeders and their male helpers, suggesting that males increasetheir investment in reproductive capability under the risk ofsperm competition. Sperm competition models also predict thatlarger testes are associated with increased siring success.Our paternity analysis supported this prediction; we found apositive relationship between testis investment by male helpersand the number of offspring they sired.     

ASPECTS OF THE BREEDING BIOLOGY OF THE FIERYNECKED NIGHTJAR

Jackson H. D. 1985. Aspects of the breeding biology of the Fierynecked Nightjar. Ostrich 56: 263–276.

A marked population of nightjars in Zimbabwe was studied intensively for four breeding seasons. This paper covers certain aspects of the breeding biology of the Fierynecked Nightjar Caprimulgus pectoralis. The male shows strong site fidelity during the breeding season (September to December), singing, feeding and breeding within an area of about 5,8 ha. There is some evidence of site defence by the male. The female shows strong mate fidelity, resulting in a pair bond for life. Egg laying starts with full moon in September and is further stimulated by the next two full moon periods. The eggs are laid directly on dense leaf litter at a site overhung by foliage. The normal clutch is two eggs (12S % are one egg) laid on successive days during the afternoon. Incubation starts with the first egg and is by the male at night and the female by day. The incubation period is 18 days. The birds respond to undue disturbance by deserting the eggs and laying a replacement clutch. The chicks usually hatch on successive afternoons; they are mobile on the first day and will move to a parent if called. Both parents feed and brood the young during twilight and moonlight; the male broods them on dark nights and the female does so by day. The species is double-brooded when time permits, the female laying again once the first brood has reached independence; she may even lay a third clutch if the second one comes to grief. There is no evidence of adults transporting eggs or young.