Evolutionary morphology of the caecilian urogenital system. I. The gonads and the fat bodies

The comparative morphology of the gonads and fat bodies of members of 17 genera and 46 species of caecilians (Amphibia: Gymnophiona) is described and analyzed. Comparison is made with the morphology of salamanders and frogs in order to elucidate evolutionary trends and relationships within the order Gymnophiona and within the class Amphibia. The structure of the testis lobes and transverse and longitudinal ducts is described based on gross dissection and histological investigation. The pattern of spermatogenesis and interstitial tissue changes are described and compared with those of other amphibians. A trend toward fusion of testis lobes is analyzed. The characteristics of the seasonal reproductive cycle of male Gymnopis m. proxima are described, and evidence for cyclic reproductive activity in other forms is presented. The morphology of the ovaries and ova is described. Size of ovary and size and number of ova is dependent on the state of maturation of the ova. Some evidence for seasonal ovum production and breeding is presented. Fat body morphology is found to be correlated with size, nutrition, and gonad condition, as in other amphibians.     

Developmental plasticity and maternal effects of reproductive characteristics in the frog,Bombina orientalis

Summary Life history theory suggests that reproductive characteristics such as ovum size and clutch size should be well buffered against vararies of the environment. However, studies which demonstrate environmental sensitivity of reproductive characteristics are increasing in number, as are studies which find that maternal effects are responsible for much of the variation in developmental and growth rates in embryonic and larval fish and amphibians. The data reported here demonstrate that the environment, in terms of temperature and food availability that a specific individual encounters during vitellogenesis, exerts a strong influence on both egg size and number. Warmer temperatures and less food decrease ovum size, while colder temperatures and less food decrease clutch size. The variation in ovum size that is induced by the environment can exert a strong influence on variation in offspring development and growth and serve as an excellent model for studies on the evolution of developmental plasticity.     

Reproductive pattern of the freshwater prawn Pseudopalaemon bouvieri (Crustacea,Palaemonidae) from hypo-osmotic shallow lakes of Corrientes (Argentina)

Pseudopalaemon bouvieri undergoes complete abbreviated development. The reproductive cycle (proportion and size of the sexually mature population and juvenile recruitment), fecundity and egg size of this prawn were examined in three subtropical shallow lakes. The reproductive cycle was seasonal; gonadal maturation occurred during the winter, ovigerous females were abundant in the spring and breeding occurred at the end of spring. Females produced small numbers of eggs (9–55) of relatively large sizes (1.0–2.1 mm). The proportion of ovigerous females with respect to the total number of females, the fecundity and egg size differed among the lakes according to the trophic state of the environments. Pseudopalaemon bouvieri has a reproductive strategy similar to other Palaemonidae species that inhabit nutrient-poor inland waters.     

Plasticity of reproductive traits responding to variation in light availability at the rosette stage of the first year in a strict biennial,Pedicularis torta,from a field on the Qinghai–Tibet Plateau,China

Light is a fundamental heterogeneous environmental factor for plants and often varies at both spatial and temporal scales. In the present study, we carried out a shading experiment to investigate the effect of variable light availability at the rosette stage of the first year in the strict biennial species Pedicularis torta (an endemic species of the Qinghai‐Tibet Plateau) on reproductive traits (i.e. total number of flowers per plant, total number of fruits per plant/total number of flowers per plant, the number of seeds per fruit, mean individual seed mass and estimation of the total number of seeds per plant, plant size at maturity in the second year and on the relationships between reproductive traits and plant size at this stage). We found that light availability affected most of the reproductive traits (except for mean individual seed mass), plant size at maturity in the second year, and the relationship between the total number of flowers per plant and plant size at this stage. Our results indicate that reproductive traits respond to changes in light availability during the rosette plant stage.     

Reproduction mode of European Bitterling (Rhodeus amarus,Bloch, 1782) determined through rapid oocyte counts and size determination using digital imaging

This study focuses on assessing the reproduction mode of an important model species for evolutionary and behavioural ecology by using digital image analysis: the European bitterling Rhodeus amarus (Bloch, 1782). Specifications of mode of reproduction were determined using oocyte size distribution, seasonal dynamics in mean oocyte diameter and total number of oocytes in ovary samples gained from April to July 2007. The rapid oocyte count was enabled by using lucia image analysis software, which also provided measurement and colour estimations of oocytes. Bitterling ovaries showed features typical for indeterminate spawners, i.e. a continuous distribution of oocyte size over the reproductive season and recruitment of new pre‐vitelogenic oocytes in the second half of the reproductive season. These results are consistent with the view that the European bitterling is a batch spawning fish with indeterminate fecundity.     

Is the reproductive potential of wild house mice regulated by extrinsic or intrinsic factors?

Abstract The regulation of reproductive performance in small mammals may be determined by extrinsic or intrinsic parameters. In a large‐scale, replicated field experiment we monitored the seasonal fluctuation in food availability and tested the effects of food addition on the reproductive performance of wild house mice (Mus domesticus) in south‐eastern Australia. Ovulation rates and litter size increased during spring and peaked in October/November. Ovulation rate was consistently higher than litter size by approximately 1.2 embryos (19%). None of the extrinsic parameters measured (food quality and quantity, mouse abundance) had an impact on reproductive performance. The addition of food did not prevent the mid summer decrease in ovulation rates nor did it alter the difference between ovulation rates and litter size. While the number of previous pregnancies did not affect reproductive performance, the age of mice did: older mice tended to have higher ovulation rates than younger mice. The effect of age‐dependent changes in ovulation rates on population growth rates of house mice seemed to be of limited importance. We conclude that the reproductive output in wild house mice is determined by ovulation rates and not by litter size. The regulation of ovulation rates through an intrinsic factor (age) seems evident but the importance of food availability and house mouse abundance for ovulation rates is low.     

Induced responses of nestling great tits reduce hen flea reproduction

The dynamics of host–parasite interactions depend to a large extent on the effect of host responses on parasite fitness. Exposure to parasites may induce behavioural or physiological responses in hosts that may reduce the subsequent survival or reproductive output of the parasite. Neonate hosts may further directly obtain immunologically active substances from their mother, for instance via milk in mammals or egg yolk in birds. However, the relative importance of maternally‐derived and self‐generated responses in inducing parasite resistance is poorly understood, especially in free‐living vertebrates. Here we investigate the complementary effect of experimentally induced maternal and neonate responses in great tit (Parus major) hosts on the reproductive success of their common ectoparasite, the hen flea (Ceratophyllus gallinae). In the laboratory we measured the number of eggs and larvae produced by individual flea females collected from host nests. In addition, the total number of larvae produced by an experimentally set number of flea females in the host's nestbox was assessed under field conditions. There was no indication of maternally‐transferred parasite resistance, since exposing the mother to fleas during the laying period did not affect the reproductive rate of fleas exploiting her offspring early or late in the nestling cycle. Independent of the maternal treatment, exposure of neonates to fleas early in the nestling period reduced the reproductive output of fleas late in the nestling cycle. The effect of the induced nestling response was seasonal, reducing flea reproduction in nests of early‐breeding hosts but not in nests of late‐breeding ones. Larvae production in the nestbox and in the laboratory was positively correlated, but under natural conditions the neonate response did not affect the size of the flea larvae population. Our results indicate induced responses as a means by which neonate avian hosts resist ectoparasites. Other factors, such as the environmental temperature and density‐dependent larval competition, may be more important in determining the size of the future parasite populations.     

Seasonal variation of egg size and number in a Daphnia pulex population

Seasonal variation of egg size and number was examined in a Daphnia pulex population inhabiting a vernal pond. In this population, size at maturity declines at midseason, probably as an adaptive response to size-selective predation by larvae of the salamander Ambystoma. The larger early season individuals produce more and larger eggs than the smaller late season individuals. Age at maturity does not vary between seasons. Laboratory experiments indicate that temperature may affect egg size, egg number and size at maturity. However, field data suggest that temperature accounts for only a small fraction of the total variation in egg size and number. Indirect measures of nutrition indicate that food limitation does not cause the seasonal decline in egg size and number. The seasonal change in reproductive traits is well correlated with changes in invertebrate and vertebrate predation. Examination of predator feeding preferences and their impact on Daphnia mortality indicate that variation of reproductive traits is most likely a complex adaptation to changing predation regimes.     

Reproductive success and individual variation in feeding frequency of House Sparrows (Passer domesticus)

Parental care is assumed to be closely associated with individual differences in reproductive success. We investigated how feeding frequencies varied among parents and how this affected the subsequent reproductive success in insular populations of House Sparrows Passer domesticus in northern Norway. Female parents fed their offspring more than male parents did, and the feeding rates were positively related to the feeding rates of the partner. A positive relationship between feeding rates and bill depth was present in females. In males, the feeding rates were negatively related to total badge size and positively related to visible badge size, after the effect of other variables had been taken into account. A non-linear convex relationship between feeding frequency and hatch day was present in males, which could reflect either the seasonal change in weather conditions or the seasonal variation in food availability. For both sexes, feeding frequencies increased with increasing brood size, but at the same time the average feeding rate per nestling decreased with increased brood sizes. Finally, our results indicate that the amount of parental investment, measured as feeding rates during the nestling stage, may have a positive long-term influence both on the number of fledglings that recruit as well as the probability that fledglings survive until recruitment.     

Genome‐wide association study for reproductive traits in a Large White pig population

Using the PorcineSNP80 BeadChip, we performed a genome‐wide association study for seven reproductive traits, including total number born, number born alive, litter birth weight, average birth weight, gestation length, age at first service and age at first farrowing, in a population of 1207 Large White pigs. In total, we detected 12 genome‐wide significant and 41 suggestive significant SNPs associated with six reproductive traits. The proportion of phenotypic variance explained by all significant SNPs for each trait ranged from 4.46% (number born alive) to 11.49% (gestation length). Among them, 29 significant SNPs were located within known QTL regions for swine reproductive traits, such as corpus luteum number, stillborn number and litter size, of which one QTL region associated with litter size contained the ALGA0098819 SNP for total number born. Subsequently, we found that 376 functional genes contained or were near these significant SNPs. Of these, 14 genes—BHLHA15, OCM2, IL1B2, GCK, SMAD2, HABP2, PAQR5, GRB10, PRELID2, DMKN, GPI, GPIHBP1, ADCY2 and ACVR2B—were considered important candidates for swine reproductive traits based on their critical roles in embryonic development, energy metabolism and growth development. Our findings contribute to the understanding of the genetic mechanisms for reproductive traits and could have a positive effect on pig breeding programs.     

Reproductive condition of Akodon montensis Thomas and Bolomys lasiurus (Lund) (Rodentia, Muridae) based on histological and histometric analyses of testes and external characteristics of gonads

The reproductive activity of two species of wild rodents was studied in the Serra do Rola‐Moça State Park (PERM), in the state of Minas Gerais, Brazil. From October 1998 to October 1999 specimens of Akodon montensis (Thomas 1913) were captured in an area of seasonal semi‐deciduous forest, while specimens of Bolomys lasiurus (Lund 1840) were captured in a cerrado area (a savannah‐like grassland). The determination of the reproductive condition was based on histological and histometric analyses of testes and external characteristics of gonads. The volume of the testes, diameter of the seminiferous tubules and depth of the seminiferous epithelium were measured. Akodon montensis showed a seasonal annual reproductive cycle divided into four stages (recrudescence, activity or proliferation, regression and rest) with the reproductive activity mainly occurring during the rainy season. In contrast, B. lasiurus presented reproductive activity during almost the whole year. For both species the volume of the testes was an effective indicator of their reproductive condition. We suggest that the histological and histometric analysis of the gonads of males, together with the observations of external characteristics of gonads of males and females, may assist in identifying the reproductive condition of these animals.     

Development of secondary sexual characters and their relationship to ontogeny and seasonal reproductive period in Hyphessobrycon igneus (Ostariophysi: Characiformes)

Sexual dimorphism in size, anal‐fin shape and coloration of Hyphessobrycon igneus, Characidae, were examined. Males were more frequent at larger body sizes, confirming body size as a sexually dimorphic trait. Anal‐fin shape and the colour of all fins were the same for females and juveniles, differing only in adult males. Likewise, only adult males had bony hooks on fin rays; larger and more sexually mature males had the most numerous and developed hooks and hooks were most developed in degree and number during peak reproductive periods. Fin hooks regressed in number and developmental degree after the reproductive period, but restarted development with the beginning of the new reproductive period without completely disappearing. Results show that bony hooks have a development and regression cycle related to reproductive seasonality.     

Maternal investment in reproduction and its consequences in leatherback turtles

Maternal investment in reproduction by oviparous non-avian reptiles is usually limited to pre-ovipositional allocations to the number and size of eggs and clutches, thus making these species good subjects for testing hypotheses of reproductive optimality models. Because leatherback turtles (Dermochelys coriacea) stand out among oviparous amniotes by having the highest clutch frequency and producing the largest mass of eggs per reproductive season, we quantified maternal investment of 146 female leatherbacks over four nesting seasons (2001–2004) and found high inter- and intra-female variation in several reproductive characteristics. Estimated clutch frequency [coefficient of variation (CV) = 31%] and clutch size (CV = 26%) varied more among females than did egg mass (CV = 9%) and hatchling mass (CV = 7%). Moreover, clutch size had an approximately threefold higher effect on clutch mass than did egg mass. These results generally support predictions of reproductive optimality models in which species that lay several, large clutches per reproductive season should exhibit low variation in egg size and instead maximize egg number (clutch frequency and/or size). The number of hatchlings emerging per nest was positively correlated with clutch size, but fraction of eggs in a clutch yielding hatchlings (emergence success) was not correlated with clutch size and varied highly among females. In addition, seasonal fecundity and seasonal hatchling production increased with the frequency and the size of clutches (in order of effect size). Our results demonstrate that female leatherbacks exhibit high phenotypic variation in reproductive traits, possibly in response to environmental variability and/or resulting from genotypic variability within the population. Furthermore, high seasonal and lifetime fecundity of leatherbacks probably reflect compensation for high and unpredictable mortality during early life history stages in this species.     

Reproductive aspects of the Atlantic angel shark Squatina dumeril

Atlantic angel sharks Squatina dumeril were collected by fishery‐dependent and independent trawls from 2002 to 2008 for reproductive analysis. Female S. dumeril have only one functional ovary (left), with an average litter size of seven pups. The reproductive cycle is at least biennial, though the seasonality of vitellogenesis could not be determined. Gestation is c. 12 months, and embryo data support a seasonal trend in reproduction, with parturition occurring in the spring months (February to June). Mature male S. dumeril have spines on the outer margins of their pectoral fins, and there is an apparent peak in gonad size in the spring. The total length at which 50% of the population is mature is 85·8 and 92·9 cm for females and males, respectively.     

Reproductive biology of Haemulon plumierii in the south‐western Atlantic Ocean's most extensive reefs: implications for fisheries management

The reproductive biology of the white grunt Haemulon plumierii was studied from 360 individuals obtained from artisanal fisheries landings in the Abrolhos Bank, Brazil, between August 2010 and March 2012. The overall sex‐ratio did not differ significantly from 1:1, although males predominated in larger size classes. β‐Binomial modelling of historical sex‐ratio data indicated that the catch rate of females has increased in recent years. Females reached maturity at a smaller total length (L_T; 214 mm) than males (235 mm L_T) and the L_T at which 50% of all individuals are mature (L₅₀) was 220 mm, corresponding to 41·5% of the maximum recorded L_T. Variation in the gonado‐somatic index and in the relative frequency of reproductive stages indicates that reproduction occurs year round, with increased activity during the austral spring and summer. Fecundity was not size dependent. The reproductive parameters provided here can support management measures focussed on seasonal closures during spawning peaks (September to November and February to March) and minimum sizes (>L₅₀) for the capture of this important artisanal fisheries resource in Abrolhos, the region with the largest and most biodiverse coralline reefs in the South Atlantic Ocean.     

Plant size influences abundance of floral visitors and biomass allocation for the cushion plant Thylacospermum caespitosum under an extreme alpine environment

Variation in size may influence the abundance of visitors and reproductive allocation for cushion plants in the extreme alpine environments. To assess effects of plant size on the abundance of main visitors and reproductive allocation in Thylacospermum caespitosum populations at two altitudes, the abundance of the visitors, visiting frequency, total number of flowers, number of fruits, number of unseeded flowers, and reproductive allocation were investigated during the period of reproductive growth. Concurrently, the effects of plant size on the visitors' contributions to fruit setting rate were assessed by a bagging experiment. Our results showed that flies (Musca domestica and Dasyphora asiatica) were the main pollinating insects of T. caespitosum, and they could obvious facilitate (p < 0.05) the fruit setting rate of this cushion plant. Seed set and floral visitation were significantly influenced (p < 0.001) by plant size. Moreover, the reproductive allocation and fruit setting rate of T. caespitosum was influenced (p < 0.001) by plant size. More biomass was allocated to reproduction in plants of greater diameter. There is an increase in reproductive success (increases of fruit number with increase in plant size) in relation to plant size. In conclusion, the extent of M. domestica and D. asiatica to facilitate the fruit setting rate mainly depended on the size of T. caespitosum. Size‐dependent reproductive allocation occurred in T. caespitosum and was the chief factor affecting the contribution of flies to fruit setting rate. These traits reflect reproductive fitness of T. caespitosum related to plant size in extreme alpine environments.     

Contrasting reproductive traits in two species of mangrove‐dwelling littorinid snails in a seasonal tropical habitat

Previous studies on the reproductive biology of littorinid snails have focused on rocky shore species, investigating how these gastropods can achieve maximal reproductive success, as well as on processes of sexual selection. This study documented differences in the reproductive traits of two mangrove‐dwelling littorinids, Littoraria ardouiniana and L. melanostoma, in Hong Kong. Reproductive activity of both species was most intense during the summer months. Mating pairs of the two species generally occurred in the tree canopies. Few false mating pairs (same sex or heterospecific pairs: <10%) were recorded, and members of both species showed size‐assortative mating. Littoraria ardouiniana had a shorter reproductive season but a higher intensity of mating and higher seasonal fecundity, than did L. melanostoma. Members of both species showed bi‐lunar periodicities of egg or larval release, synchronized with spring tides. Fecundity showed a strong positive relationship with body size in L. ardouiniana, but not in L. melanostoma. Females of L. ardouiniana released entire broods of larvae in a single brief event, whereas females of L. melanostoma released fewer eggs over 1–8 d. Release of larvae in L. ardouiniana involved a series of short bursts and was much faster than the trickle release of eggs in L. melanostoma. The contrasting reproductive traits in these two species represent different strategies to optimize reproductive success in mangrove habitats.     

Reproductive strategy of Leporinus piau (Fowler, 1941), a neotropical freshwater fish from the semi‐arid region of Brazil

This study concerns the reproductive strategy of the freshwater fish, Leporinus piau, native to the Brazilian semi‐arid region. The hypothesis of this work predicts that the species would spawn during the rainy season, presenting a seasonal reproductive strategy. The study investigated the structure of total body length and body mass, sex ratio, body size at first sexual maturity, developmental stages of gonads, fecundity, gonadosomatic index (GSI), condition factor (K) and the spawning season of L. piau. Twelve monthly samplings (mean n = 18) were carried out in 2009 and environmental parameters such as rainfall, temperature, pH, electrical conductivity and concentration of dissolved oxygen of water were registered. The sampled population of L. piau (n = 211) showed a slight predominance of males (55%), but with larger and heavier females. First sexual maturation in males occurred earlier (16.5 cm total length) than in females (20.5 cm total length). The species is a total spawner with an average fecundity of 55 000 mature oocytes. Rainfall (225 ± 7.2 mm) and concentration of dissolved oxygen (8.5 ± 0.2 mg L^?1) acted as influential factors during the spawning season. The monthly GSI shows that spawning starts when the rainy season is at its maximum and ends synchronously (females r = 0.91; males r = 0.89) with the end of the rainy season. L. piau presents a seasonal reproductive strategy with high reproductive output, providing a population increase and exploitation of the environmental resources.     

Reproductive seasonality in captive wild ruminants: implications for biogeographical adaptation,photoperiodic control,and life history

Many ruminant species show seasonal patterns of reproduction. Causes for this are widely debated, and include adaptations to seasonal availability of resources (with cues either from body condition in more tropical, or from photoperiodism in higher latitude habitats) and/or defence strategies against predators. Conclusions so far are limited to datasets with less than 30 species. Here, we use a dataset on 110 wild ruminant species kept in captivity in temperate‐zone zoos to describe their reproductive patterns quantitatively [determining the birth peak breadth (BPB) as the number of days in which 80% of all births occur]; then we link this pattern to various biological characteristics [latitude of origin, mother‐young‐relationship (hider/follower), proportion of grass in the natural diet (grazer/browser), sexual size dimorphism/mating system], and compare it with reports for free‐ranging animals. When comparing taxonomic subgroups, variance in BPB is highly correlated to the minimum, but not the maximum BPB, suggesting that a high BPB (i.e. an aseasonal reproductive pattern) is the plesiomorphic character in ruminants. Globally, latitude of natural origin is highly correlated to the BPB observed in captivity, supporting an overruling impact of photoperiodism on ruminant reproduction. Feeding type has no additional influence; the hider/follower dichotomy, associated with the anti‐predator strategy of ‘swamping’, has additional influence in the subset of African species only. Sexual size dimorphism and mating system are marginally associated with the BPB, potentially indicating a facilitation of polygamy under seasonal conditions. The difference in the calculated Julian date of conception between captive populations and that reported for free‐ranging ones corresponds to the one expected if absolute day length was the main trigger in highly seasonal species: calculated day length at the time of conception between free‐ranging and captive populations followed a y = x relationship. Only 11 species (all originating from lower latitudes) were considered to change their reproductive pattern distinctively between the wild and captivity, with 10 becoming less seasonal (but not aseasonal) in human care, indicating that seasonality observed in the wild was partly resource‐associated. Only one species (Antidorcas marsupialis) became more seasonal in captivity, presumably because resource availability in the wild overrules the innate photoperiodic response. Reproductive seasonality explains additional variance in the body mass–gestation period relationship, with more seasonal species having shorter gestation periods for their body size. We conclude that photoperiodism, and in particular absolute day length, are genetically fixed triggers for reproduction that may be malleable to some extent by body condition, and that plasticity in gestation length is an important facilitator that may partly explain the success of ruminant radiation to high latitudes. Evidence for an anti‐predator strategy involving seasonal reproduction is limited to African species. Reproductive seasonality following rainfall patterns may not be an adaptation to give birth in periods of high resource availability but an adaptation to allow conception only at times of good body condition.     

The considerable adult size variability in wood feeders is optimal

1. The life history of wood feeders was modelled in order to explain the multiseasonality of development and the great variability of adult size in this group. 2. The model was parameterised with experimental bioenergetic and reproductive data for the xylem feeder Aredolpona rubra (Coleoptera: Cerambycidae). 3. The length of the developmental period, which together with food quality directly determines adult size and indirectly determines the number of eggs laid, was optimised. 4. The results show that multi‐seasonal larval development maximises fitness under conditions of low food quality, relatively low predation pressure, and the presence of hostile periods during the year. 5. The variability of the number of seasons needed to complete development within a wood‐feeder population is a consequence of development time optimisation and the unavoidable extension of the egg‐laying period. These insects have an evolutionary dilemma: to eclose late in a given season at smaller size, bringing about low egg production and low offspring value, or to grow bigger to the next season, jeopardising their lives. 6. The results of the model predict wood‐feeder developmental patterns that depend on the tree tissue inhabited.