Marker-assisted determination of the relationship between body size and reproductive success and consequences for evaluation of adaptive life histories

We tested for differences in the predicted optimal ages at first maturity in brook charr ( Salvelinus fontinalis ) in Freshwater River, Newfoundland, when life-history data were collated based on the marker-assisted estimation of the relationship between body size and reproductive success rather than using fecundity as a surrogate for reproductive success. Jointly with capture–recapture data to estimate the growth and survival costs of reproduction, we found that weak relationships between body size and reproductive success generate selection against delayed maturation. This finding would not have held for females if the relationship between body size and fecundity had been used as a surrogate for the relationship between body size and reproductive success. This shows that predictions of optimal life histories can be qualitatively changed when using molecular markers to directly evaluate age- and/or size-specific effects of body size on reproductive success.     

State-dependent male mating tactics in the grey seal: the importance of body size

The purpose of this study was to examine the importance of bodysize and body composition as determinants of conditional matingtactics exhibited in male grey seals. We combined behavioralobservations with measures of energy expenditure and successon 42 known-age individuals during the breeding seasons of 1997–2001at Sable Island, Canada. Males with a large body mass arrivedat the breeding grounds with more body fat and body energy andcatabolized less body protein than smaller males. Males consumed1.9 ± 0.2 MJ day^–1, and those with a smaller percentageof body fat had higher rates of food energy intake. The amountof body energy on arrival was positively correlated with theduration of the breeding period. Males that exhibited the primarymating tactic of consortship were heavier, had absolutely morebody fat and body energy, and sustained breeding longer thanthose males that did not exhibit the primary tactic. Amongstall males, body mass showed a quadratic relationship with thenumber of female consorts mated and the estimated number ofpups sired. Thus, intermediate-sized males mated with the mostconsorts and achieved the highest success. Intermediate bodysize may be optimal during breeding due to greater agility inmale combat. Body size was an important determinant of matingtactics used by male grey seals. A large body size providedan energetic advantage of greater endurance while an intermediatebody size may provide greater competitive ability in acquiringconsortships.     

The influence of horn and body size on the reproductive behavior of the horned rainbow scarab beetlePhanaeus difformis (Coleoptera: scarabaeidae)

The reproductive behavior of horned rainbow scarab beetles,Phanaeus difformis, was studied to determine the influence of morphological traits on intersexual and intrasexual interactions.Phanaeus difformis is a sexually dimorphic dung beetle in which males possess much larger horns than females, and males can be grouped into major and minor male morph categories based on horn size. Male-female pairs cooperated in nest construction and provisioning. In the laboratory, males of both morphs assisted females and were equally successful at copulating. However, in the field larger individuals had a pairing advantage due to greater success in intrasexual competition. Some males used an alternative mating tactic which involved sneaking copulations with paired females. In most cases the sneak male was smaller than the paired male.     

The influence of body size on immediate reproductive success in Dugesia gonocephala (Tricladida,Paludicola)

The individual reproductive output of the stream-dwelling flatworm Dugesia gonocephala was investigated. Various measures of reproductive success were related to body size. (I) For the first 30 days in the laboratory small individuals produced no cocoons, individuals of intermediate size produced unfertilized cocoons and large individuals usually produced fertilized cocoons. (II) In individuals that produced a cocoon, no correlation was found between the number of cocoons produced in one month and body size. (III) Large individuals, however, produced larger cocoons. This was not due to the fact that unfertilized cocoons were smaller. (IV) Large cocoons tended to contain more young. (V) The average size of young hatching from large cocoons was larger. (VI) Large individuals produced their first cocoon soon after their arrival in the laboratory and seemed to have a higher chance of producing a fertilized first cocoon. (VII) A trade-off existed between producing many small versus few large young.     

Initial reproductive investment and parental body size in Cryptocercus punctulatus (Dictyoptera: Cryptocercidae)

Abstract. Females of the subsocial woodroach Cryptocercus punctulatus Scudder generally have a single oviposition period during which they produce one to four oöthecae. Monogamous pairs and their recently deposited oöthecae were collected in the field, and measured, weighed and analysed for nitrogen in the laboratory. Females put 9.8 ± 2.4% (mean ± SD) of their dry weight and 11.6 ±0.5% of their body nitrogen into oöthecae. The total nitrogen and dry weight of the brood were positively related to the post-oviposition total nitrogen and dry weight of the mother. A female's investment, on a per nymph basis, averaged 0.06% of her post-oviposition dry weight. Females may be able to recover up to 58.7% of the nitrogen invested into a brood by consuming the egg cases after hatch. Overall, there was little variation in the width of head capsules of adults in this species, and this parameter was not significantly different between the sexes. Females were heavier than males (dry weight) (P = 0.06). Within pairs, the weights and nitrogen contents of males and females were positively correlated, probably because they feed together in the same log for nearly a year prior to reproduction.     

Ecological correlates of body size and egg size in parasitic Ascothoracida and Rhizocephala (Crustacea)

In order to identify key factors in the evolution of life history traits in Ascothoracida and Rhizocephala (two groups of crustacean parastes of invertebrates), comparative analyses were performed using phylogenetically independent contrasts. Among 59 ascothoracidan species, latitude correlated positively with body size, whereas there was no relationship between water depth and body size. Body size also correlated strongly with egg size; however, once corrected for body size, egg size was not related to either latitude or water depth. Among 91 rhizocephalan species, neither latitude nor water depth correlated with body size. However, host species of larger sizes harboured larger species of rhizocephalan parasites. Egg size of rhizocephalans did not correlate with body size, and was not influenced by either latitude or water depth. The patterns observed in this study show both differences from an similarities to those reported for other groups of crustacean parasites, and suggest that adaptations to similar selective pressures are not always identical among distantly-related taxa.     

The life history legacy of evolutionary body size change in carnivores

We estimate the body sizes of direct ancestors of extant carnivores, and examine selected aspects of life history as a function not only of species' current size, but also of recent changes in size. Carnivore species that have undergone marked recent evolutionary size change show life history characteristics typically associated with species closer to the ancestral body size. Thus, phyletic giants tend to mature earlier and have larger litters of smaller offspring at shorter intervals than do species of the same body size that are not phyletic giants. Phyletic dwarfs, by contrast, have slower life histories than nondwarf species of the same body size. We discuss two possible mechanisms for the legacy of recent size change: lag (in which life history variables cannot evolve as quickly as body size, leading to species having the 'wrong' life history for their body size) and body size optimization (in which life history and hence body size evolve in response to changes in energy availability); at present, we cannot distinguish between these alternatives. Our finding that recent body size changes help explain residual variation around life history allometries shows that a more dynamic view of character change enables comparative studies to make more precise predictions about species traits in the context of their evolutionary background.     

Life History Studies on the Genus Trillium (Liliaceae) I. Reproductive Biology of Four Japanese Species

Abstract Four Japanese Trillium (Liliaceae) species which are representative perennial herbs of the temperate deciduous forests— Trillium kamtschaticum (2x), T. apetalon (4x), T. tschonoskii (4x) and T. smallii (6x)—were studied for their reproductive characteristics, e.g., patterns of reproductive resource allocation and reproductive output.
In spite of the differences in ploidy levels (from 2x to 6x), all four species showed very similar reproductive traits. It became evident that in response to the increase in reproductive allocation to total reproductive organs (RA), the number of seeds produced per plant (P_N) clearly increased. This trend is well in accord with the relationship found in four North American species and also several temperate woodland perennial herbs which occur in closed, stable and predictable environments, and possess typical xenogamous breeding systems. Although there is no conspicuous trend between ploidy levels and P_N, one of the significant differences noted in this study was in seed weight, which ranged from 2.93 mg in diploids, to 3.42–3.45 mg in tetraploids, and to 4.47 mg in hexaploids.     

Life history patterns in female moose (Alces alces): the relationship between age,body size,fecundity and environmental conditions

I examined the relationship between age, body size and fecundity in 833 female moose (Alces alces) from 14 populations in Sweden sampled during 1989–1992. Data on population density, food availability and climatic conditions were also collected for each population. Age and body mass were both significantly positively related to fecundity, measured as ovulation rate, among female moose. The relationship between the probability of ovulation and body mass was dependent on age with (1) a higher body mass needed in younger females for attaining a given fecundity, and (2) body mass having a stronger effect on fecundity in yearling (1.5 year) than in older (2.5 year) females. Thus, a 40 kg increase in yearling body mass resulted in a 42% increase in the probability of ovulation as compared to a 6% increase in older females. The lower reproductive effort per unit body mass, and the relatively stronger association between fecundity and body mass in young female moose compared to older ones, is likely to primarily represent a mechanism that trades off early maturation against further growth, indicating a higher cost of reproduction in young animals. In addition to age and body mass, population identity explained a significant amount of the individual variation in fecundity, showing that the relationship between body mass and fecundity was variable among populations. This variation was in turn related to the environment, in terms of climatic conditions forcing female moose living in relatively harsh/more seasonal climatic conditions to attain a 22% higher body mass to achive the same probability of multiple ovulation (twinning) as females living in climatically milder/less seasonal environments. The results suggests that the lower fecundity per unit body mass in female moose living in climatically harsh/more seasonal environments may be an adaptive response to lower rates of juvenile survival, compared to females experiencing relatively milder/less seasonal climatic conditions.     

Subsocial behaviour in Oniticellus cinctus (Coleoptera, Scarabaeidae): effect of the brood on parental care and oviposition

ABSTRACT. In Oniticellus cinctus (F.) the nest chambers each contain about twenty brood balls. Females enlarge the brood balls during the egg and larval stages and remain in the chamber for the whole period of brood development (1 month); they then make a new nest after 1 week. The presence of the brood releases parental care and ensures that the mother remains in the nest: she repairs defects in the brood balls and the nest, and expels other O. cinctus females. A new ball is formed around a naked O. cinctus larva, but larvae of other species are killed. In addition, the brood inhibits oviposition: removal (or addition) of brood balls stimulates (or inhibits) egg laying. In inhibited ovarioles, existing follicles are resorbed and production of new ones ceases. Control of clutch size by the brood is an adaptation to the nest structure and life history of O. cinctus. It may have an important role in the reproductive strategy of other insects with parental care.     

Life history evolution in cichlids 2: directional evolution of the trade-off between egg number and egg size

The negative relationship between offspring number and offspring size provides a classic example of the role of trade-offs in life history theory. However, the evolutionary transitions in egg size and clutch size that have produced this negative relationship are still largely unknown. Since body size may affect both of these traits, it would be helpful to understand how evolutionary changes in body size may have facilitated or constrained shifts in clutch and egg size. By using comparative methods with a database of life histories and a phylogeny of 222 genera of cichlid fishes, we investigated the order of evolutionary transitions in these traits in relation to each other. We found that the ancestral large-bodied cichlids first increased egg size, followed by a decrease in both body size and clutch size resulting in the common current combination of a small-bodied cichlid with a small clutch of large eggs. Furthermore, lineages that deviated from the negative relationship between clutch and egg size underwent different transitions in these traits according to their body size (large bodied genera have moved towards the large clutch/small egg end of the continuum and small bodied genera towards the small clutch/large egg end of the continuum) to reach the negative relationship between clutch size and egg size. Our results show that body size is highly important in shaping the negative relationship between clutch size and egg size.     

Sexual behaviour and evolution of sexual dimorphism in body size in Jaera (Isopoda Asellota)

Individuals of the genus Jaera do not mate at random. In the species from the Mediterranean group, J. italica and. J. nordmanni, large males and medium sized females are at an advantage and their sizes are positively assorted. These effects are attributable to sexual competition between males. In the Ponlo-caspian species J. istri, no advantage of large males exists, but sexual selection could be the cause for a long passive phase prior to copulation and for normalizing selection upon female size at pairing. In the Atlantic species, J. albifrons, no selection can be ascertained.
Differential mating success in males appears as one of the causes of the evolution of sexual dimorphism in body size, which makes males larger, of equal size, or smaller than females according to the species. The reason for this reversal in dimorphism seems to differ in the two sexes. Sexual selection provides an explanation for the evolution of male size, while the interspecific changes in female length are more likely due to ecological factors.     

Experimental analysis of an early life‐history stage: avian predation selects for larger body size of hatchling turtles

One common life‐history pattern involves an elevated rate and nonrandom distribution of neonatal mortality. However, the mechanisms causing this pattern and the specific traits that confer a survival benefit are not always evident. We conducted a manipulative field experiment using red‐eared slider turtles to test the hypothesis that diurnal avian predators are a primary cause of size‐specific neonatal mortality. Body size was a significant predictor of recapturing hatchlings alive and of finding hatchlings dead under natural conditions, but was unimportant when diurnal predators were excluded from the field site. Overall recapture rates also more than doubled when predators were excluded compared to natural conditions (72.4 vs. 34.9%). We conclude that birds are an important cause of size‐specific mortality of recently emerged hatchling turtles and that ‘bigger is better’ in this system, which has important implications for life‐history evolution in organisms that experience size‐specific neonatal mortality.     

Correlated evolution of colour pattern and body size in polymorphic pygmy grasshoppers, Tetrix undulata

Theory posits that selection on functionally interrelated characters will promote physical and genetic integration resulting in evolution of favourable trait-value combinations. The pygmy grasshopper Tetrix undulata (Orthoptera: Tetrigidae) displays a genetically encoded polymorphism for colour pattern. Colour morphs differ in several traits, including behaviours, thermal biology and body size. To examine if these size differences may reflect phenotypic plasticity of growth and development in response to temperature we used a split brood-design and reared hatchlings from mothers belonging to different morphs in different thermal environments (warm or cold) until maturity. We found that time to maturity was longer in the cold compared with the warm treatment. In the warm (but not in the cold) treatment time to maturity also varied among individuals born to mothers belonging to different colour morphs. Although low temperature and long development time are normally accompanied by increased body size in ectotherms, our results revealed no difference in size at maturity between individuals reared in the two temperature treatments. There was also an increase (not a decrease) in adult body size with shortened time to maturity across families within each treatment. Taken together, this suggests that body size is canalized against environmental perturbations, and that early maturation does not necessarily trade off against a size-mediated decrease in fecundity. Heritability of body size was moderate in magnitude. Moreover, body size at maturity varied among individuals belonging to different morphs and was influenced also by maternal colour morph, suggesting that a genetic correlation exists between colour pattern and body size. These findings suggest that different characters have evolved in concert and that the various colour morphs represent different evolutionary strategies, i.e., alternative peaks in a multi-modal adaptive landscape.     

Life history and reproductive capacity of Gammarus fossarum and G. roeseli (Crustacea: Amphipoda) under naturally fluctuating water temperatures: a simulation study

SUMMARY 1. Mathematical functions developed in long‐term laboratory experiments at different constant temperatures were combined with daily water temperatures for 1991–93 in eight Austrian streams and rivers to simulate the complex life histories and reproductive capacities of two freshwater amphipods: Gammarus fossarum and G. roeseli. The functions describe brood development times, hatching success, times taken to reach sexual maturity, growth, and fecundity. The sex ratio was assumed to be 0.5 and an autumn–winter reproductive resting period was based on observations of six river populations. Simulations included summer‐cold mountain streams, summer‐warm lowland rivers, watercourses fed by groundwater or influenced by heated effluents, and varying amplitudes of change within each year. 2. A fortran 77 computer program calculated growth from birth to sexual maturity of first‐generation females born on the first day of each calendar month in 1991, and the numbers of offspring successfully released from the maternal broodpouch in successive broods. At the 1991–93 regimes of temperature, individual G. fossarum released 127–208 offspring and G. roeseli released 120–169 in seven or eight successive broods during life spans of less than 2 years in six rivers. Life spans extended into a third year in the relatively cool River Salzach (mean temperature 7.5 °C). They were not completed in the very cold River Steyr (mean 5.6, range 2.5–7.9 °C), where G. fossarum produced five broods (totalling 120 offspring) and G. roeseli only two broods (totalling 28 offspring) in the 3‐year period. Except in the Steyr, some offspring grew rapidly to maturity and produced several second‐generation broods during the simulation period; in the warmest rivers some third‐generation broods were also produced. Birth dates, early or late in the year, influenced the subsequent production of broods and young, depending on temperature regimes in particular rivers. Total numbers of offspring produced by the second and third generations represent the theoretical reproductive capacities of G. fossarum and G. roeseli. Minimum and maximum estimates mostly ranged from 100 to 17 300, were larger for G. fossarum except in the warmest river (March), where temperatures rose above 20 °C for 56–78 days in summer, and largest (maximum 37 600) in the River Voeckla heated by discharge from a power‐station (mean 11.5 °C). Results from the simulations agree with preliminary assessments of relative abundances for G. fossarum and G. roeseli in several of the study rivers, but in some one or both species appear to be absent. On a wider scale, the present study confirms that G. fossarum is potentially more successful than G. roeseli in cool rivers but indicates that neither species is likely to maintain viable populations in cold rivers strongly influenced by snow and ice‐melt. 3. The potential impacts of future river warming by increases of 1, 2 and 3 °C, due to climate change, vary according to river site, date of fertilisation, the extent of temperature increase, and the species of Gammarus. For Austrian rivers with mean temperatures in the range c. 7–10 °C, future warming would result in modest changes in the life histories and reproductive capacities of both G. fossarum and G. roeseli; the former would find improved temperature conditions in watercourses that are currently very cold throughout the year, and both would find warm rivers less tolerable. 4. The high potential reproductive capacity of gammarids, with rapid production of numerous successive broods when sexual maturity is finally achieved, indicates adaptation to high mortality during the relatively long period of growth to sexual maturity, and provides scope for an opportunistic strategy of emigration from centres of population abundance to colonise new territory when conditions are favourable. Rapid expansion of populations is desirable to combat the effects of environmental catastrophes, both frequent and short‐term floods and droughts, and more long‐term climatic changes that have occurred several times in glacial–interglacial periods during the current Ice Age.     

Genetic estimates of annual reproductive success in male brown bears: the effects of body size, age, internal relatedness and population density

1. We studied male yearly reproductive success (YRS) and its determinants (phenotypic characteristics, age, population density) in two Scandinavian brown bear populations, using molecular techniques to determine paternity. 2. We found a significant difference in male YRS between the study areas, with lower YRS in the south than in the north. 3. In general, older and larger males had higher YRS. Older males may be more experienced in competition for reproduction (male dominance). Large body size is of direct benefit in male-male competition and of advantage in endurance competition for the access to females. 4. Age was relatively more important for YRS in the north and body size was more important in the south, due perhaps to differences in male age structure due to illegal killing. A single old male dominated the reproduction in the north during the study, which resulted most probably in the relatively higher importance of age in the north. In the south, with a more even male age structure, no single male was able to dominate, probably resulting in a more intense competition among males, with body size as the deciding factor. 5. Male YRS was correlated positively with population density. This may be related to the structure of the expanding bear population, with female densities declining towards the population edge. 6. Internal relatedness, a measure of genetic heterozygosity, was correlated negatively with YRS, suggesting that outbred individuals have a higher YRS. Individual heterozygosity at key or many loci may reflect male physical qualities and condition-sensitive traits, which may benefit males directly in contest or in sperm competition.     

Trade-offs between reproductive and somatic (storage) investments in animals: a comparative test of the Van Noordwijk and De Jong model

Classical life-history theory predicts ‘trade-offs’ between reproductive and somatic investments. However, empirical studies have shown that intraspecific phenotypic correlations between these two resource investments are often positive or nonsignificant, rather than negative as predicted. The model of Van Noordwijk and De Jong (1986) was proposed to explain these unexpected results. According to their model, positive correlations between reproductive and somatic investments will result if individual variation in resource acquisition exceeds that of resource allocation, whereas negative correlations will result if individual variation in resource allocation exceeds that of resource acquisition. To test this model, I used body storage/condition as an index of somatic investment because it is usually strongly related to level of resource acquisition. I predicted that laboratory studies should more often show negative correlations between reproductive and somatic investments than field studies, because individual variation in resource acquisition is expected to be lower in controlled laboratory environments than in variable natural environments. A literature review revealed that correlations between somatic (storage) investment and reproductive investment (estimated as clutch/litter mass, number of offspring per clutch/litter, or number of clutches/litters) among conspecific breeding female animals are more often positive (15 species) or nonsignificant (17 species) than negative (6 species). Moreover, as expected, five of six negative correlations were observed in laboratory studies, whereas 13 of 15 positive correlations were observed in field studies. It is concluded that future empirical and theoretical work on life histories should consider individual variation in both resource acquisition and allocation and the interaction between the two. This revised version was published online in July 2006 with corrections to the Cover Date.