A Review for Life-history Traits Variation in Frogs Especially for Anurans in China

Environmental variation can promote differentiation in life-history traits in species of anurans. Increased environmental stress usually results in larger age at sexual maturity, older mean age, longer longevity, slower growth, larger body size, and a shift in reproductive allocation from offspring quantity to quality, and a stronger trade-off between offspring size and number. However, previous studies have suggested that there are inconsistent geographical variations in life-history traits among anuran species in China. Hence, we here review the intraspecific patterns and differences in life-history traits(i.e., egg size, clutch size, testes size, sperm length, age at sexual maturity, longevity, body size and sexual size dimorphism) among different populations within species along geographical gradients for anurans in China in recent years. We also provide future directions for studying difference in sperm performance between longer and shorter sperm within a species through transplant experiments and the relationships between metabolic rate and brain size and life-history.     

Insular shifts and trade-offs in life-history traits in pond frogs in the Zhoushan Archipelago, China

Island and mainland populations of animal species often differ strikingly in life-history traits such as clutch size, egg size, total reproductive effort and body size. However, despite widespread recognition of insular shifts in these life-history traits in birds, mammals and reptiles, there have been no reports of such life-history shifts in amphibians. Furthermore, most studies have focused on one specific life-history trait without explicit consideration of coordinated evolution among these intimately linked life-history traits, and thus the relationships among these traits are poorly studied. Here we provide the first evidence of insular shifts and trade-offs in a coordinated suite of life-history traits for an amphibian species, the pond frog Rana nigromaculata . Life-history data were collected from eight islands in the Zhoushan Archipelago and neighboring mainland China. We found consistent, significant shifts in all life-history traits between mainland and island populations. Island populations had smaller clutch sizes, larger egg sizes, larger female body size and invested less in total reproductive effort than mainland populations. Significant negative relationships were found between egg size and clutch size and between egg size and total reproductive effort among frog populations after controlling for the effects of body size. Therefore, decreased reproductive effort and clutch size, larger egg size and body size in pond frogs on islands were selected through trade-offs as an overall life-history strategy. Our findings contribute to the formation of a broad, repeatable ecological generality for insular shifts in life-history traits across a range of terrestrial vertebrate taxa.     

Reproductive strategies and body shape in the European pond turtle (Emys orbicularis) from contrasting habitats in Italy

Comparisons within and among populations offer important insights into variation in life-history traits and possible adaptive patterns to environmental conditions. We present the results of observed differences in body size, body shape and patterns of reproduction in four separate populations of the European pond turtle Emys orbicularis in central and southern Italy – coastal ( n =3) and mountainous ( n =1) sites and pond ( n =2) and canal ( n= 2) habitats – to determine whether phenotypic plasticity affects reproductive output. Although we did not find any significant latitudinal variation in body size, we observed significant differences in body shape between canal (rounded body shape) and pond (elongated body shape) systems and smaller size with rounded shape in the mountainous population. Reproductive output is similar among populations (median=5 eggs per clutch), whereas reproductive investment (relative clutch mass to maternal body mass) is higher in the mountain population (one clutch per year) than in coastal populations (two clutches per year), suggesting differential trade-offs between geographic locality, elevation and habitat type. Turtle shell shape and geographic location together affect reproductive output in E. orbicularis in Italy.     

Reproductive ageing and sexual selection on male body size in a wild population of antler flies (Protopiophila litigata)

Little is known about the importance of trade-offs between ageing and other life history traits, or the effects of ageing on sexual selection, particularly in wild populations suffering high extrinsic mortality rates. Life history theory suggests that trade-offs between reproduction and somatic maintenance may constrain individuals with higher initial reproductive rates to deteriorate more rapidly, resulting in reduced sexual selection strength. However, this trade-off may be masked by increased condition dependence of reproductive effort in older individuals. We tested for this trade-off in males in a wild population of antler flies (Protopiophila litigata). High mating rate was associated with reduced longevity, as a result of increased short-term mortality risk or accelerated ageing in traits affecting viability. In contrast, large body size was associated with accelerated ageing in traits affecting mating success, resulting in reduced sexual selection for large body size. Thus, ageing can affect sexual selection and evolution in wild populations.     

Life-history strategies of Acrossocheilus fasciatus (Barbinae, Cyprinidae) in the Huishui Stream of the Qingyi watershed, China

Life-history traits of Acrossocheilus fasciatus were examined using 384 specimens collected monthly during May 2009 and April 2010 in the Huishui Stream of the Qingyi watershed, China. Using scales for age determination, female and male fish comprised five and four age groups, respectively. The monthly changes in marginal increment ratio suggested that annuli on scales were formed during March through May. Total lengths back-calculated significantly increased with age for both sexes and varied significantly between the two sexes at each age. The fact that females had larger body size and grew faster than males indicated the sexual size dimorphism for this species. Both sexes got their 50% maturity at age 3, when females and males were 105.3 and 112.1?mm total length, respectively. Based on the monthly changes in the gonado-somatic index and egg-development process, fish spawned from April through August. Absolute fecundity ranged from 295 to 3,573 eggs per fish and increased significantly with age. But relative fecundity, ranging from 11.77 to 69.96?eggs/g, was not significantly different among age groups. Compared with the life-history traits of an upstream population in the Puxi Stream (a headwater stream within this study watershed), the downstream population of A. fasciatus in the Huishui Stream (a 4th-order stream) exhibits larger body size, faster somatic growth, later sexual maturity, and lower reproductive investment. These variations in life-history strategies between the two populations could perhaps be explained by the spatial heterogeneity in habitat environment along the upstream–downstream gradient in this watershed.     

Reproductive cycles and reproductive strategies among populations of the Rose‐bellied Lizard Sceloporus variabilis (Squamata: Phrynosomatidae) from central Mexico

Species with wide distribution, generally show variations in life history characteristics, which can be attributed to environmental causes. In this study, we analyzed the reproductive cycle and reproductive characteristics from three populations (Atlapexco, San Pablo Tetlapayac, and Santa Catarina) of the lizard Sceloporus variabilis from central Mexico. The specific goal of this study was to evaluate life history characteristics such as reproductive period extent, SVL (snout‐vent length) at sexual maturity, clutch size, egg mass and volume, and RCM (relative clutch mass). The San Pablo Tetlapayac population showed a larger clutch size, RCM, egg mass, and a smaller SVL, body mass and reproductive period (January‐September), as well as egg volume than the Atlapexco and Santa Catarina populations. Reproductive cycle and reproductive characteristics were more similar between the Atlapexco and Santa Catarina populations. Differences found in the population of San Pablo Tetlapayac with respect to the Atlapexco and Santa Catarina populations could be attributed to environmental variations where lizard populations occur. Differences in the reproductive period and reproductive characteristics in each population could be the result of both historical (phylogenetic; e.g., reproductive mode) and nonhistorical (environmental; e.g., temperature, food availability) causes. This study showed that populations of the same species are under different selection pressures, and these affect the reproductive characteristics of populations. Our results also indicate that long‐term and targeted studies on predation, use and selection of food, are needed to determine the causes of these variations in populations of S. variabilis.     

内蒙古中东部草原不同生境克氏针茅（Stipa krylovii Roshev.）种群的形态差异分析

对内蒙古中东部草原分布的克氏针茅进行了种群内和种群间的形态差异分析。结果表明：（1）种群内不同的形态性状存在不同程度的差异;（2）种群间生殖枝中的一部分形态性状以及营养枝高度和营养枝干重均存在较大差异;（3）每穗小花数、每穗籽粒数、每穗小穗数、生殖枝高、穗干重、生殖枝干重、每穗种子重、种子重/生殖枝重比等性状的变化趋势相同,均表现为随生境条件变差而增加,即在生殖上投入增加以增大适合度;（4）主成分分析表明穗长、种子重/生殖枝重比、每穗小花数、每穗籽粒数、生殖枝高、基盘长、第二芒柱长、营养枝高、千粒重、芒针长是不同种群差异的主要指标。这些形态性状的差异可以看作克氏针茅对不同生境的适应性表现。     

Variation of reproductive traits in a population of the lizard Lacerta vivipara

Variation in reproductive traits (sexual maturity, clutch size, clutch weight, mean egg mass, newborn weight) was studied during a four year period in a population of the live-bearing lizard Lacerta vivipara . Sexual maturity was associated with attaining a minimum body size. Clutch size increased with female body length and litter weight increased with clutch size. A major component of the within year variation in these reproductive traits was attributable to female size. Analysis of successive clutches in individual females indicated that a significant fraction of the variation in litter size, adjusted for female length, was due to consistent differences between individuals. Newborn weight varied within and among litters, but no relations between hatchling mass or mean egg mass in a litter and other traits were detected.
Size-adjusted reproductive performances remained constant during the course of this study, even though environmental conditions (weather factors, food availability) varied annually. Observed among year variations in reproductive characteristics were attributable to differences in the body size distributions of the adult females.     

Old world versus new world: life-history alterations in a successful invader introduced across Europe

We examined differences in pumpkinseed (Lepomis gibbosus) life-history traits between native North American and introduced European populations, and tested three life-history predictions related to the effect of temperature, growth, waterbody size, and the presence/absence of predators on native and non-native populations. Pumpkinseed populations exhibit more ‘opportunistic’ traits (earlier maturity, smaller size at maturity, and higher reproductive allocation) in their introduced European range than those in their native range. Predictions of life-history traits were improved when indicators of juvenile growth rate (mean length at age 2), waterbody size (surface area), and thermal regime (air temperature degree-days above 10 °C) were incorporated into models along with continental location, but European pumpkinseed populations exhibit more opportunistic life-history traits than North American populations even when these factors are accounted for. Native pumpkinseed in waterbodies containing piscivores mature later and at a larger size, and have lower gonadosomatic indices than those in waterbodies lacking piscivores, whereas there is no significant difference in the same three life-history traits between European waterbodies containing or lacking piscivores. Because congeneric competitors of the pumpkinseed are absent from Europe, the apparent absence of a predator life-history effect there could also be due to the absence of the major sunfish competitors. In either case, the evolution and maintenance of more opportunistic traits in European pumpkinseed can likely be attributed to enemy release, and this may explain the successful establishment and spread of pumpkinseed in many parts of Europe.     

Consistency in the life history traits of four invasive pseudorasbora parva populations in Southern England

Life history plasticity can be a strong predictor of the establishment and invasion success of introduced fishes. Here, the life-history traits of four P. parva populations in adjacent ponds in Southern England were measured throughout 2013 to determine the timing and length of their reproductive season and the extent of trait plasticity. The relative abundance of the populations (as catch per unit effort) was similar, with low variability in their traits relating to reproductive effort and somatic growth. All the populations were male dominated. Both sexes matured at small body sizes, with fish as small as 30 mm being mature in both sexes, with the age at maturity for both sexes being age 1+ years. The peak spawning period, characterised by female fish investing heavily in reproduction when their gonado-somatic index (GSI) values were highest and declined thereafter, occurred in May and June, and after a mean of 212 ± 24 degree-days >12°C, and with water temperatures being 13.2–14.6°C. There were no further peaks in GSI, despite the species being considered to have prolonged spawning periods across spring and summer. These results also suggest that whilst the expression of life-history traits of invasive fishes often vary with density, they can be highly consistent between populations at similar abundances and locations.     

Population divergence along lines of genetic variance and covariance in the invasive plant Lythrum salicaria in eastern North America

Evolution during biological invasion may occur over contemporary timescales, but the rate of evolutionary change may be inhibited by a lack of standing genetic variation for ecologically relevant traits and by fitness trade-offs among them. The extent to which these genetic constraints limit the evolution of local adaptation during biological invasion has rarely been examined. To investigate genetic constraints on life-history traits, we measured standing genetic variance and covariance in 20 populations of the invasive plant purple loosestrife (Lythrum salicaria) sampled along a latitudinal climatic gradient in eastern North America and grown under uniform conditions in a glasshouse. Genetic variances within and among populations were significant for all traits; however, strong intercorrelations among measurements of seedling growth rate, time to reproductive maturity and adult size suggested that fitness trade-offs have constrained population divergence. Evidence to support this hypothesis was obtained from the genetic variance-covariance matrix (G) and the matrix of (co)variance among population means (D), which were 79.8% (95% C.I. 77.7-82.9%) similar. These results suggest that population divergence during invasive spread of L. salicaria in eastern North America has been constrained by strong genetic correlations among life-history traits, despite large amounts of standing genetic variation for individual traits.     

SPATIAL AND TEMPORAL VARIATION IN SELECTION ON CORRELATED LIFE-HISTORY TRAITS AND PLANT SIZE IN CHAMAECRISTA FASCICULATA

Intraspecific studies of selection on multiple traits of a plant's life history provide insight as to how the composite life history of an organism evolves. Current understanding of selection on plant life-history traits is deficient in three important areas: 1) the effects of selection through correlated traits, 2) the effects of selection on a trait throughout the plant's lifetime, and 3) spatial and temporal variation in selection on plant life-history traits among populations and years. This study documents spatial and temporal variation in selection on three life-history and two morphological traits for two natural populations of Chamaecrista fasciculata, a native summer annual. Life-history and morphological traits (date of seedling emergence, size at establishment, size prior to reproduction, date of initial flowering, and date of initial fruit maturation) varied significantly between sites and/or years. Selection on traits varied either spatially, between sites and among transects within one site, or temporally, between years. In addition, life-history traits were phenotypically correlated among themselves and with morphological traits; correlations were generally constant over time and space. Indirect selection caused changes in means and variances in traits not under direct selection, but which were correlated with traits under selection. Selection on date of emergence varied in direction and magnitude among different life-cycle stages, while selection on other traits varied only in magnitude among life stages of the plant. This study documents the complexity of the selective process and the importance of considering multiple life stages and traits when studying the evolution of life-history traits.     

Differences in life-history traits in two clonal strains of the self-fertilizing fish, <Emphasis Type="BoldItalic">Rivulus marmoratus</Emphasis>

Synopsis We compared life-history traits such as fecundity, sex ratio, reproductive cycle, age at sexual maturity, embryonic period, egg size, early growth and morphology in two clonal strains (PAN-RS and DAN) of the mangrove killifish, Rivulus marmoratus, under constant rearing conditions. We found a positive relationship between growth and reproductive effort. Fecundity was significantly higher in the PAN-RS strain than in the DAN strain. The sex ratio was significantly different, with DAN producing more primary males than PAN-RS. Spawning and ovulation cycle did not clearly differ between the strains. PAN-RS showed a significantly higher growth rate than DAN from 0 to 100 days after hatching, however, age at sexual maturity, embryonic period, egg size, and morphometric and meristic characteristics (vertebral and fin-ray counts) did not differ between the two strains. The high fecundity of PAN-RS may provide an increased chance of offspring survival, while the attainment of sexual maturity at a smaller size in DAN may allow them to invest earlier in reproduction to increase breeding success. Variations in the life-history traits of PAN-RS and DAN may be adaptive strategies for life in their natural habitat, which consists of mangrove estuaries with a highly variable environment.     

Density effects on life-history traits of an island lizard population

Long-term monitoring of life-history traits and the effects of density upon them were studied in an island population of the lizardEumeces okadae. Although life-history traits such as clutch size, egg size and the proportion of mature reproductive females varied little over 7 years in the intact population, manipulation of density to simulate decreased population density enhanced juvenile growth rate, age at first reproduction, frequency of female reproduction and size-specific clutch mass. In particular, the proportion of mature females reproducing annually increased almost 10 times from 5.6% to 53.8% after the removal of some lizards. However, body size at first reproduction and egg size were almost identical under both high and low density conditions. This study suggests that there were strong density-dependent effects on several life-history traits and thatE. okadae attained a density close to the carrying capacity of the environment.     

Phenotypic plasticity in reproductive traits induced by food availability in a lacertid lizard, Takydromus septentrionalis

Understanding the proximate determinants of phenotypic variations in life-history traits can provide powerful insights into a species' life-history strategies. I experimentally manipulated availability of food (high vs low) to examine plasticity in the reproductive traits of northern grass lizards, Takydromus septentrionalis (Lacertidae), from eastern China. Food availability significantly affected reproductive frequency and thereby seasonal reproductive output, but had little effect on reproductive output per clutch. Low-food females postponed reproduction and produced less clutches in the reproductive season than did high-food females. After producing their second clutches, low-food females were in lower body condition than the high-food counterparts. By the end of the experiment, however, all females exhibited similar body condition. Clutch size and clutch mass differed between the first and second clutches but not between the treatments. Egg size and phenotypic traits of hatchlings (body size, morphology and locomotor performance) in T. septentrionalis did not vary significantly from first to second clutches nor between the two treatments. These results support optimal egg size (offspring) theory. Female T. septentrionali s "decide" whether or not to reproduce largely based on current energy intake; lowered feeding rates thus delay oviposition and reduce reproductive frequency. In contrast, clutch size, egg size and relative clutch mass remain unchanged.     

Evolution of pleiotropic alleles for maturation and size as a consequence of predation

Understanding life-history evolution requires knowledge about genetic interactions, physiological mechanisms and the nature of selection. For platyfish, Xiphophorus maculatus, extensive information is available about genetic and physiological mechanisms influencing life-history traits. In particular, alleles at the pituitary locus have large and antagonistic effects on age and size at sexual maturation. To examine how predation affects the evolution of these antagonistic traits, I examined pituitary allele evolution in experimental populations differing in predation risk. A smaller size, earlier maturation allele increased in frequency when predators were absent, while a larger size, later maturation allele increased in frequency when predators were present. Thus, predation favours alleles for larger size, at the cost of later maturation and reproduction. These findings are interesting for several reasons. First, predation is often predicted to favour early reproduction at smaller sizes. Second, few studies have shown how selection acts on alleles that affect age and size at sexual maturation. Finally, many studies assume that trade-offs between these life-history traits result from antagonistic pleiotropy, with alleles that positively affect one trait negatively affecting another, yet this is rarely known. This study unequivocally demonstrates that genetically based trade-offs affect life-history evolution in platyfish.     

Sexual and lifetime selection on body size in a marine fish: the importance of life-history trade-offs

Many field measurements of viability and sexual selection on body size indicate that large size is favoured. However, life-history theory predicts that body size may be optimized and that patterns of selection may often be stabilizing rather than directional. One reason for this discrepancy may be that field estimates of selection tend to focus on limited components of fitness and may not fully measure life-history trade-offs. We use an 8-year, demographic field study to examine both sexual selection and lifetime selection on body size of a coral reef fish (the bicolour damselfish, Stegastes partitus). Selection via reproductive success of adults was very strong (standardized selection differential=1.04). However, this effect was balanced by trade-offs between large adult size and reduced cumulative survival during the juvenile phase. When we measured lifetime fitness (net reproductive rate), selection was strongly stabilizing and only weakly directional, consistent with predictions from life-history theory.     

The evolution of life-history traits in parasitic and free-living platyhelminthes: a new perspective

Parasite life histories have been assumed to be shaped by their particular mode of existence. To test this hypothesis, we investigate the relationships between life-history traits of free-living and parasitic platyhelminthes. Using phylogenetically independent contrasts we examine patterns of interspecific covariation in adult size, progeny volume, daily fecundity, total reproductive capacity, age at first reproduction and longevity. The correlations obtained indicate a similar causal chain of life history variations for free-living and parasitic platyhelminthes. These results suggest that increased longevity favours delayed reproduction. Furthermore, growth pattern determines adult body size and age at maturity. For platyhelminthes, whether free-living or parasitic, the total reproductive capacity is found to be directly determined by the size of the worm. Within this group the parasitic way of life does not seem to influence the basic patterns of life history evolution. Received: 20 September 1997 / Accepted: 1 March 1998     

Reaction norms of size and age at maturity of Pomacea canaliculata (Gastropoda: Ampullariidae) under a gradient of food deprivation

Pomacea canaliculata, an apple snail native to South America,has become a serious pest of aquatic crops and a promoter ofecosystem changes in natural wetlands worldwide. Its successas an invader has been attributed to its great phenotypic plasticityin life-history traits. Our aims were to determine the reactionnorms of size and age at maturity under a gradient of food deprivation.Full sibling experimental snails were reared in isolation fromhatching and maintained until maturity under seven differentlevels of relative food deprivation based on size-specific ingestionrates. To detect the onset of sexual activity of experimentalsnails, fully mature virgin snails reared in the laboratorywere used as consorts. The reaction norms for age and size atmaturity of P. canaliculata showed marked sexual dimorphism.Shell length was the main component of variation in the malereaction norms for both copulation and egg-laying by femaleconsorts, whereas age was the main component of variation forfemales. Irrespective of the intensity of food deprivation,males mature at the same age at the expense of size, since sizeis apparently irrelevant in the access to females and male fitnesscan be maximized through fast maturation. In contrast, a minimumsize is required for females to reach maturity, perhaps as aresult of their higher reproductive costs. The highly dimorphicreaction norms lead to an increasing lag between male and femalematurity as deprivation increases; in temperate regions, malesborn early in the reproductive season would mature in the sameseason irrespective of food availability, while most femaleswould have to overwinter before attaining sexual maturity inunproductive habitats or those dominated by unpalatable macrophytes.The great life-history plasticity reported in invaded areascould be a heritage from populations in the native range. (Received 10 August 2007; accepted 30 July 2008)     

Variation in Body Shape across Species and Populations in a Radiation of Diaptomid Copepods

Inter and intra-population variation in morphological traits, such as body size and shape, provides important insights into the ecological importance of individual natural populations. The radiation of Diaptomid species (~400 species) has apparently produced little morphological differentiation other than those in secondary sexual characteristics, suggesting sexual, rather than ecological, selection has driven speciation. This evolutionary history suggests that species, and conspecific populations, would be ecologically redundant but recent work found contrasting ecosystem effects among both species and populations. This study provides the first quantification of shape variation among species, populations, and/or sexes (beyond taxonomic illustrations and body size measurements) to gain insight into the ecological differentiation of Diaptomids. Here we quantify the shape of five Diaptomid species (family Diaptomidae) from four populations each, using morphometric landmarks on the prosome, urosome, and antennae. We partition morphological variation among species, populations, and sexes, and test for phenotype-by-environment correlations to reveal possible functional consequences of shape variation. We found that intraspecific variation was 18-35% as large as interspecific variation across all measured traits. Interspecific variation in body size and relative antennae length, the two traits showing significant sexual dimorphism, were correlated with lake size and geographic location suggesting some niche differentiation between species. Observed relationships between intraspecific morphological variation and the environment suggest that divergent selection in contrasting lakes might contribute to shape differences among local populations, but confirming this requires further analyses. Our results show that although Diaptomid species differ in their reproductive traits, they also differ in other morphological traits that might indicate ecological differences among species and populations.