Allocation in reproduction is not tailored to the probable number of matings in common toad (<Emphasis Type="Italic">Bufo bufo</Emphasis>) males

The theory of life history evolution assumes trade-offs between competing fitness traits such as reproduction, somatic growth, and maintenance. One prediction of this theory is that if large individuals have a higher reproductive success, small/young individuals should invest less in reproduction and allocate more resources in growth than large/old individuals. We tested this prediction using the common toad (Bufo bufo), a species where mating success of males is positively related to their body size. We measured testes mass, soma mass, and sperm stock size in males of varying sizes that were either (1) re-hibernated at the start of the breeding season, (2) kept without females throughout the breeding season, or (3) repeatedly provided with gravid females. In the latter group, we also estimated fertilization success and readiness to re-mate. Contrary to our predictions, the relationship between testes mass and soma mass was isometric, sperm stock size relative to testes mass was unrelated to male size, fertilization success was not higher in matings with larger males, and smaller males were not less likely to engage in repeated matings than larger males. These results consistently suggest that smaller males did not invest less in reproduction to be able to allocate more in growth than larger males. Causes for this unexpected result may include relatively low year-to-year survival, unpredictable between-year variation in the strength of sexual selection and low return rates of lowered reproductive investment.     

Reproduction of asteroids from Balsfjorden,Northern Norway: Analyses of lipids in the gonads of Ctenodiscus crispatus,Asterias lincki and Pteraster militaris

Seasonal lipid analyses were performed on the gonads and residual bodies of Ctenodiscus crispatus (Retzius) and Asterias lincki (Müller & Troschel), collected from Balsfjorden, northern Norway during March 1978–February 1980. The ovaries of both species had high percentages of total lipid (in the region of 60% of the dry weight) that were relatively constant throughout the season. Testes from both species had in the region of 20% of their dry weight as lipid, again with little seasonal variation. The majority of the total lipid in the ovaries of C. crispatus was accounted for by alkyldiacylglycerols and there was little seasonal variation in lipid class composition. Phospholipid accounted for the bulk of the lipid in the testes of C. crispatus, where alkyldiacylglycerols were present but in minor amounts. A slight degree of seasonal variation occurred in the composition of testis lipid in that maximal percentages of triacylglycerols and alkyldiacylglycerols occurred in January–April. The ovaries of A. lincki contained both triacylglycerols and alkyldiacylglycerols as major neutral lipids. Significant seasonal variation occurred in lipid class composition in the ovaries in that neutral lipids showed transient minima in February–April. The testes of A. lincki had substantial amounts of both triacylglycerols and alkyldiacylglycerols: maximal percentages of these lipids occurred in January–March. Analyses of gonads of near-mature Pteraster militaris (O. F. Müller) revealed that the ovaries were rich in lipid that was predominantly alkyldiacylglycerols, whereas the testes had much lower levels of lipid accounted for mainly by phospholipids. The residual bodies of all three asteroids had significant but small amounts of alkyldiacylglycerols in their lipids. A. lincki had triacylglycerols as a major lipid class in its body lipids, whereas in C. crispatus the major body lipid was phospholipid. Little seasonal variation was seen in the composition of the residual body lipid of either C. crispatus or A. lincki. The results are consistent with an aseasonal reproductive pattern in C. crispatus, although a main breeding activity occurred in mid-winter. A. lincki is a more seasonal breeder, but also had maximal activity in winter. Irrespective of their breeding habits, all three asteroids produced eggs rich in alkyldiacylglycerols. A major role for this lipid in the development of the large lecithotrophic eggs of asteroids is indicated.     

Larval alarm pheromones as a potential control for invasive cane toads (Bufo marinus) in tropical Australia

Novel approaches to control invasive species are urgently needed. Cane toads (Bufo marinus) are large, highly toxic anurans that are spreading rapidly through tropical Australia. Injured toad larvae produce an alarm pheromone that elicits rapid avoidance by conspecifics but not by frog larvae. Experiments in outdoor ponds show that repeated exposure to the pheromone reduced toad tadpole survival rates (by >50%) and body mass at metamorphosis (by 20%). The alarm pheromone did not induce tadpoles to seek shelter, but accelerated ontogenetic differentiation. Perhaps reflecting mortality of weaker individuals during larval life, growth rates post-metamorphosis were higher in animals emerging from the pheromone exposure treatment than from the control treatment. Nonetheless, body size differentials established at metamorphosis persisted through the first 8 days of post-metamorphic life. We will need substantial additional research before evaluating whether the alarm pheromone provides a way to reduce cane toad recruitment in nature, but our field trials are encouraging in this respect.     

Testes mass in megachiropteran bats varies in accordance with sperm competition theory

Sperm competition is a widespread phenomenon influencing a range of characters, including investment in gonadal tissue. Conspecific proximity is one factor which can influence the risk of sperm competition and hence testicular investment, and decreased confidence of paternity may be one cost of group living. Aspects of female biology may also influence spermatogenic investment and sperm morphology. This study examines the associations between relative testes mass and roost-group size across 17 species of Megachiroptera. Associations between breeding season duration and investment in spermatogenesis are also examined, as are associations between female reproductive tract dimensions and testes mass and dimensions of spermatozoa across all bats. Relative testes mass was significantly positively associated with roost-group size at a species level and after appropriate phylogenetic control (pairwise comparisons and comparison of independent contrasts). There were no significant relationships between breeding season duration and relative testes mass. Across all bats, neither testes mass nor sperm length were significantly related to dimensions of the female tract. The results are discussed in the context of sperm competition. Received: 7 January 1998 / Accepted after revision: 8 August 1998     

Growth rate and the cost of calling activity in male carpenter frogs,Rana virgatipes

Summary Energy stress during the breeding season and relationships between calling activity and growth were investigated in male carpenter frogs, Rana virgatipes. This species has a prolonged breeding season of up to three months in Southern New Jersey. Monthly collections made in 1985 revealed that both dry mass and percent body lipid decreased throughout the breeding season but sharply increased at the end of the breeding season. Observations of free-living males showed that small males were more likely to gain mass than large males during the breeding season. All males gained mass at higher rates after the breeding season. A simultaneous record of calling activity and mass change was obtained for 42 males. Males called on 95% of nights, indicating that they rarely ceased their reproductive activities. Small males tended to have low calling efforts and high growth rates. When the effect of initial mass was removed, growth rate was negatively correlated with calling effort among small males. This is the first demonstration of a direct tradeoff between a reproductive activity and growth in an anuran.     

Song amplitude and body size in birds

Bird song is a sexually selected multidimensional signal. A fundamental question regarding the evolution of sexually selected signals is what information they convey and how their honesty is maintained. Song amplitude is a performance-related signal trait that varies considerably between individuals, but this signal dimension has been neglected in past studies. I found that median song amplitude in male nightingales (Luscinia megarhynchos) and zebra finches (Taeniopygia guttata) did not vary significantly with body size or residual body mass. In contrast, I found a significant negative correlation between body size (and also residual mass) and the maximum song amplitude during interactive singing in nightingales. However, the function of these more subtle differences in song amplitude remains to be investigated. By and large, the results of this study suggest that mean song amplitude is unlikely to indicate a bird’s body size or current condition (measured as residual mass).     

Copula duration and testes size in the yellow dung fly,Scathophaga stercoraria (L.) : the effects of diet,body size,and mating history

Summary Seasonal variation in mean hind tibia length and mean testes length is investigated in the yellow dung fly, Scathophaga stercoraria (L.). There is a cycle in mean hind tibia length and mean testes length over a season. The body size curve peaks later than the testes length curve, showing that there is no fixed relationship between the two variables. The causes of variation in testes size and its influence on copula duration are experimentally examined. Increasing the number of Drosophila eaten per day leads to increased mature testes length. Males with larger testes copulate for longer than males with smaller ones, and smaller males copulate for longer than do larger males. While testes shrivel with successive copulas, copula duration remains constant. The more females a male is prepared to copulate with in a day (up to five), the longer he copulates with each. The shrinkage of the testes of males collected throughout a day suggests that males copulate with an average of 4 females per day. The costs of sperm production are thus shown to have a significant influence on the copula duration.Offprint requests to: P.I. Ward at the second address     

Spatial and temporal variation in growth rates and maturity in the Indo-Pacific squid <Emphasis Type="Italic">Sepioteuthis lessoniana</Emphasis> (Cephalopoda: Loliginidae)

Age, growth and maturity parameters are described for the Indo-Pacific squid Sepioteuthis lessoniana both temporally and spatially (equatorial, tropical and subtropical). Tropical squid that grew through periods of warming water temperatures grew 9% faster than squid that grew through periods of cool water temperatures. The tropical spring-hatched and equatorial squid had similar growth rates (3.24, 3.18 g/day) and these were significantly faster than the tropical summer/autumn hatched squid (2.89 g/day). The oldest squid aged was 224 days, but the majority of individuals were <200 days. Subtropical squid were larger, older and matured later than equatorial and tropical spring-hatched squid. The mean weight of subtropical squid aged between 100 and 150 days was >400 g, 85% greater than tropical and equatorial squid. Geographical differences revealed that subtropical mature winter females and males had mean ages >150 days, respectively 17% and 23% older than their tropical mature winter counterparts. Temporal differences in age at maturity were also evident with tropical winter females and males having a mean age of ~140 days, respectively 41% and 25% older than their summer counterparts. Cooler subtropical and winter tropical squid had the heaviest gonads (>15 g ovaries, >1.5 g testes) compared to summer tropical and equatorial squid. However, relative gonad investment (GSI) values of the cooler squid were significantly lower with cool subtropical and winter tropical females having GSI values <3, which was about half the value of the warmer water females. This study revealed considerable plasticity in the size-at-age of this species. The tropical population had growth parameters that fluctuated between an equatorial strategy (fast growth, small body size, and small gonads) and a subtropical strategy (large body size, slower growth, and large gonads) depending on season.     

On the possible contribution of mixed species flocks to species richness in neotropical avifaunas

Summary Mixed species foraging flocks are a dominant component of the infra-structure of avian communities in neotropical forests. In Amazonia, these flocks consist of pairs of 10–20 species, many of which are permanently associated with mixed flocks. At least half of these flocking species maintain territories that correspond exactly to the flock home range. Small individuals that participate as permanent members of the flocks must adopt the large home range of the larger nucleus species. Therefore, the densities of smaller species are dependent on the availability and density of flocks rather than the availability of food resources. Single pairs of 4 small flocking species with individual body masses of 8 g occupied exclusive territories of 8–12 ha. These were the same exact territories that were defended by at least 6 other flocking species with individual body masses of up to 37 g. Because of their attachment to flocks with large territories, small species are expected to under-utilize available food resources. The under-utilization of food resources is expected to allow smaller species to coexist with greater niche overlap resulting in increased species richness. This hypothesis was tested by quantifying foraging niche in terms of foraging height, foraging maneuver, and prey substrate; and using these values in addition to body mass and bill size (length, depth and width) to determine relative niche overlap between large versus small species pairs.Smaller species had greater foraging overlap than large flocking species and particularly the three smallest species of the genus Myrmotherula; longipennis, axillaris and menetriesii had very high overlap (average foraging niche overlap for the 3 species=0.83±0.12 compared with 0.12±0.19 for all flocking species), similar body sizes (body masses differing by no more then 8%) and similar bill morphologies (maximum ratio in length=1.08, width=1.07, and depth=1.06). These results are consistent with the hypothesis that small species participating in Amazonian mixed flocks can coexist with greater niche overlap because their density is flock dependent rather than resource dependent.     

Sperm traits in the quacking frog (<Emphasis Type="Italic">Crinia georgiana</Emphasis>), a species with plastic alternative mating tactics

In species where males use alternative reproductive tactics and male phenotypes are confronted with different risks of sperm competition, theory predicts that between-male-type differences in sperm expenditure may evolve. In the frog Crinia georgiana big males can monopolize females, whereas small males often engage in polyandrous matings. Consequently, big males may experience a lower risk of sperm competition than do small males. We tested if the predictions from theoretical models can be applied to the mating system of C. georgiana. Our results showed that small males do not have larger testes relative to their body size compared to their larger counterparts and that the efficiency with which sperm number, size, motility, and longevity are produced by the testes does not differ between small and large males in the predicted way. These results are not in alignment with predictions from a loaded raffle model of sperm competition on sperm expenditure in males with alternative phenotypes. The plasticity in mating tactics used by C. georgiana males and a high intraseasonal variation in male densities may have prevented the evolution of enhanced sperm performance in smaller males. A fair raffle in the sperm competition game played by C. georgiana males could also explain the observed patterns in sperm traits. Future investigations determining the parameters responsible for the deviation from theoretical predictions in this system will test the degree to which current theoretical models can indeed be applied to species with plastic reproductive tactics.     

Effects of Oil‐Palm Plantations on Diversity of Tropical Anurans

Agriculturally altered vegetation, especially oil‐palm plantations, is rapidly increasing in Southeast Asia. Low species diversity is associated with this commodity, but data on anuran diversity in oil‐palm plantations are lacking. We investigated how anuran biological diversity differs between forest and oil‐palm plantation, and whether observed differences in biological diversity of these areas is linked to specific environmental factors. We hypothesized that biological diversity is lower in plantations and that plantations support a larger proportion of disturbance‐tolerant species than forest. We compared species richness, abundance, and community composition between plantation and forest areas and between site types within plantation and forest (forest stream vs. plantation stream, forest riparian vs. plantation riparian, forest terrestrial vs. plantation terrestrial). Not all measures of biological diversity differed between oil‐palm plantations and secondary forest sites. Anuran community composition, however, differed greatly between forest and plantation, and communities of anurans in plantations contained species that prosper in disturbed areas. Although plantations supported large numbers of breeding anurans, we concluded the community consisted of common species that were of little conservation concern (commonly found species include Fejervarya limnocharis, Microhyla heymonsi, and Hylarana erythrea). We believe that with a number of management interventions, oil‐palm plantations can provide habitat for species that dwell in secondary forests. Efectos de las Plantaciones de Palma de Aceite sobre la Diversidad de Anuros Tropicales Faruk et al.     

Delayed breeding in the cooperatively breeding Florida scrub-jay (Aphelocoma coerulescens): inhibition or the absence of stimulation?

To determine whether fundamental differences exist in the reproductive physiology of breeder and nonbreeder Florida scrub-jays (Aphelocoma coerulescens), we compared plasma levels of testosterone (T) and luteinizing hormone (LH) in males, and estradiol (E₂) and LH in females. Although male breeders had higher overall T and larger testes, nonbreeders’ T paralleled that of breeders, and their testes were more than an order of magnitude larger than regressed testes. Breeder and nonbreeder males had equivalent baseline LH, and equivalent changes in LH following a gonadotropin-releasing hormone (cGnRH-I) challenge. The T, LH and GnRH challenge data indicate that nonbreeder males have functional hypothalamo-pituitary-gonadal (HPG) axes. We found no hormonal evidence of inbreeding suppression in males: nonbreeders that did not live with their mothers and those that did had similar T. Male nonbreeders that were exposed to E₂-implanted females had higher T than did controls, suggesting that the lack of within-pair stimulation is a key factor in whether an individual delays breeding. Female nonbreeders had E₂ titres equal to or higher than breeders and neither basal LH nor LH following GnRH challenge differed by breeding status. Nonbreeders’ ovarian follicles were smaller than breeders’, but were larger than they would be during the non-breeding season. These data suggest that nonbreeders were primed for breeding and were simply waiting for an opportunity or a required stimulus. Female nonbreeders that lived in a territory with an unrelated male breeder had significantly higher E₂ than those that remained with their fathers. Similarly, nonbreeders that were captured away from their home territories had elevated E₂. However, nonbreeders that lived with their fathers had E₂ that was equivalent to breeding females, suggesting that inbreeding avoidance may not be the primary factor leading to delayed breeding in females. Received: 13 June 1995 /Accepted after revision: 27 April 1996     

The significance of male-male aggression for yearling dispersal in roe deer (Capreolus capreolus)

The significance of adult buck aggression as a proximate factor driving yearling male dispersal in roe deer, Capreolus Capreolus, was studied in Sweden during the dispersal season (April–June) in 1988–1991. In roe there is a well established positive correlation between yearling antler size and dispersal tendency. I tested the prediction that yearlings with large antlers will experience more agonism from adult bucks than those with smaller antlers. Additionally, I studied the relationship between antler size and level of sexual maturity, by recording date of velvet shedding, and sampling testes weights from culled yearlings during the breeding season. The amount of agonism experienced by yearlings was positively correlated with the size of their antlers during April and May, whereas in June no such relationship was found. Aggression towards all yearlings tended to increase as the season progressed. A positive correlation between antler size and testes weight was evident, and the larger the antlers of yearlings the earlier the velvet shedding occured. These results indicate: firstly, that adult buck agonism is an important proximate factor underlying yearling male dispersal; and secondly, that agonism is mainly directed towards the sexually most mature yearlings. This, in turn, suggests mate competition to be the ultimate cause of male-male agonism, and thus male dispersal in this species.     

Fluctuating sexual dimorphism and differential hibernation by sex in a primate, the gray mouse lemur (Microcebus murinus)

The aim of this study was to investigate reproductive strategies and their consequences in gray mouse lemurs (Microcebus murinus), small solitary nocturnal primates endemic to Madagascar. Previous reports of sexual dimorphism in favor of males and females, respectively, a high potential for sperm competition and pheromonal suppression of mating activity among captive males, led us to investigate mechanisms of intrasexual competition in a wild population. Based on 3 years of mark-recapture data, we demonstrate that sexual dimorphism in this species fluctuated annually as a result of independent changes in male and female body mass. Male body mass increased significantly prior to the short annual mating season. Because their testes increased by 100% in the same period and because their canines are not larger than those of females, we suggest that large male size may be advantageous in searching for estrous females and in enabling them to sustain periods of short-term torpor. In contrast to reports from captive colonies, we found no evidence for two morphologically distinct classes of males. Finally, we also show that most adult males are active throughout the cool dry season that precedes the mating season, whereas most adult females hibernate for several months. This is in contrast to other solitary hibernating mammals, where males typically emerge 1–2 weeks before females. Thus, this first extended field study of M.␣murinus clarified previous conflicting reports on sexual dimorphism and male reproductive strategies in this primitive primate by showing that their apparent deviation from predictions of sexual selection theory is brought about by specific environmental conditions which result in sex-specific life history tactics not previously described for mammals. A general conclusion is that sexual selection can operate more strongly on males without resulting in sexual dimorphism because of independent selection on the same traits in females. Received: 6 July 1997 / Accepted after revision: 28 March 1998     

Body size preferences in the pot-bellied seahorse <Emphasis Type="Italic">Hippocampus abdominalis:</Emphasis> choosy males and indiscriminate females

Male seahorses (genus Hippocampus) provide all post-fertilization parental care, yet despite high levels of paternal investment, these species have long been thought to have conventional sex roles, with female mate choice and male–male competition. Recent studies of the pot-bellied seahorse (Hippocampus abdominalis) have shown that sex-role reversal occurs in high-density female-biased populations, indicating that male mating preferences may lead to sexual selection on females in this species. Egg size, egg number, and offspring size all correlate positively with female body size in Hippocampus, and by choosing large mating partners, male seahorses may increase their reproductive success. While male brood size is also positively correlated with body size, small H. abdominalis males can carry exceptionally large broods, suggesting that the fecundity benefits of female preference for large partners may be limited. We investigated the importance of body size in reproductive decisions of H. abdominalis, presenting focal individuals of both sexes with potential mating partners of different sizes. Mating preferences were quantified in terms of time spent courting each potential partner. Male seahorses were highly active throughout the mate-choice trials and showed a clear behavioral preference for large partners, while females showed significantly lower levels of activity and equivocal mating preferences. The strong male preferences for large females demonstrated here suggest that sexual selection may act strongly on female body size in wild populations of H. abdominalis, consistent with predictions on the importance of female body size for reproductive output in this species. An erratum to this article can be found at     

Reproduction in <Emphasis Type="Italic">Heteroteuthis dispar</Emphasis> (Rüppell, 1844) (Mollusca: Cephalopoda): a sepiolid reproductive adaptation to an oceanic lifestyle

Small cephalopods of the genus Heteroteuthis are the most pelagic members in the family Sepiolidae. This study examines the reproductive biology of Heteroteuthis dispar (Rüppell, 1844), the first such study on any member of the genus, based on 46 specimens (27 females and 19 males) collected during the Mar-Eco cruise in the North Atlantic in the region of the Mid-Atlantic Ridge in 2004, and compares it with reproductive features in the less pelagic members of the family. The unusually large spermatophores of the males have a very small ejaculatory apparatus and cement body, relative to the size of the sperm mass. Females first mate when they are still maturing: a large sperm mass (up to 3.4% of the female body mass), consisting of one to several spermatangia, was found in an internal seminal receptacle of the majority of the females examined regardless of their maturity state. The seminal receptacle has a unique form and position in this species. The receptacle is a thin-walled sac at the posterior end of the visceral mass that is an outpocketing of, and opens into, the visceropericardial coelom. Spermatangia and sperm from the spermatangia apparently enter into the visceropericardial coelom (which is mostly occupied by the ovary) from the seminal receptacle indicating that ova are fertilised internally, a strategy unknown for decapodiform cephalopods (squid and cuttlefish), but present in most octopods. Fecundity of Heteroteuthis dispar (1,100–1,300 oocytes) is much higher than in other sepiolids whereas the egg size (mean max. length ∼1.6 mm) is the smallest within the family. Spawning is continuous (sensu Rocha et al. in Biol Rev 76:291–304, 2001). These and other reproductive traits are discussed as being adaptations to an oceanic lifestyle.     

Varying levels of female promiscuity in four <Emphasis Type="Italic">Apodemus</Emphasis> mice species

Sexual selection in most vertebrates is based on the evolution of fitness optimization strategies such as multiple-male mating (MMM). Several ecological correlates of MMM have been identified in bird and fish populations; however, only few studies have documented the effects of environmental change on promiscuity in mammals. In this study, the 127 pregnant females from four central European and ecologically diverse species of field mice (genus Apodemus) were studied to assess the role of ecological factors that may have shaped the evolution of particular mating systems. MMM was found in all analyzed species: in Apodemus uralensis and Apodemus flavicollis, up to two males could be identified as the fathers of a particular litter, while three males sired 9.1% of analyzed litters of Apodemus sylvaticus and 20.6% of Apodemus agrarius. Furthermore, there were obvious differences between species in relative testes size and the proportion of multiple sired litters during those seasons when the opportunity for multiple mating was high. The species with the smallest testes and the least promiscuous was A. uralensis (only 43.5% of multiple sired litters), while the species with the biggest testes and the most promiscuous was A. agrarius (69.2%). MMM was significantly associated with higher litter size in A. flavicollis, and the probability of MMM strongly increased with season in A. agrarius and with abundance in A. uralensis. These results indicate that ecological factors are associated with MMM rates in Apodemus field mice and more research is needed to fully understand the evolution of mating strategies at different levels of biological resolution.     

Testes mass and reproductive mode of minnows

Most comparative analyses of relative testes mass find that testes are larger in species in which more sperm competition is predicted (multiple males mate with individual females). I tested for differences in adjusted testes mass (for body mass) by spawning mode and by sexual size dimorphism in a comparative analysis of 37 minnow species. No significant differences were found for testes mass by spawning mode or sexual size dimorphism. These results imply a lack of response to selection on testes size from sperm competition in minnow species. Possible explanations for the lack of the expected relationship between testes mass and mating systems in minnows are presented. Received: 8 November 1999 / Received in revised form: 27 January 2000 / Accepted: 13 February 2000     

Social system and reproduction of helpers in a cooperatively breeding cichlid fish (Julidochromis ornatus) in Lake Tanganyika: field observations and parentage analyses

It is suggested that some fish of the genus Julidochromis, substrate-brooding Tanganyikan cichlids with biparental care, breed cooperatively with helpers. We studied the social system of J. ornatus in the wild and analysed genetic parentage using microsatellites. Within the studied population three patterns of social system were identified: monogamous pairs (61%, 44 of 72 groups), pairs with helpers (29%, 21), and polygamous harems with helpers (controlled by either a large female or large male owner; 10%, 7). In cooperatively breeding groups, the number of helpers at each nest ranged from 1 to 6 (median 1), and male helpers were more numerous than female helpers. In both sexes, the body size was different among individuals of different social status (harem owner > breeder > helper). Helpers and harem owners of both sexes exhibited brood defence although its frequency was low. The molecular analysis revealed that (1) the helpers were mostly unrelated to dominant breeders, (2) many helpers of both sexes contributed genetically to the next generations, (3) male helpers had high siring success (41% of young in total), and (4) large young unrelated to group members were detected at 30% of observed nests, which may be due to breeder (or helper) replacements and immigration of young. We discuss the implications of these results for understanding the complex social system of this species, especially the low reproductive skew in comparison with other cooperatively breeding cichlids.