Smaller Adult Body Size Research Articles

Effects of individual condition and habitat quality on natal dispersal behaviour in a small rodent

1. Individuals should benefit from settling in high-quality habitats, but dispersers born under favourable conditions have a better physical condition and should therefore be more successful at settling in high-quality habitats. 2. We tested these predictions with root voles (Microtus oeconomus) by a manipulation of individual condition through litter-size enlargement and reduction during lactation combined with a manipulation of habitat quality through degradation of the vegetation cover. We accurately monitored movements of 149 juveniles during a settlement and breeding period of 3 months. 3. The litter size treatment had long-lasting effects on body size, life-history traits and home range size, but did not influence dispersal behaviour. 4. Different stages of dispersal were influenced by habitat quality. In low-quality patches, females dispersed earlier, spent more time prospecting their environment before settling, and settlers had a smaller adult body size than in high-quality patches. Preference and competition for high-quality patches is likely adaptive as it increased fitness both in terms of survival and reproduction. 5. We found no interactive effect of individual condition and habitat quality on natal dispersal and habitat selection. 6. These findings suggest that immediate conditions are more important determinants of dispersal decisions than conditions experienced early in life.

Developmental constraints on the evolution of wing‐body allometry in Manduca sexta

Artificial selection on body size in Manduca sexta produced genetic strains with large and small body sizes. The wing-body allometries of these strains differed significantly from the wild type. Selection on small body size led to a change in the scaling of wing and body size without changing the allometry: the wings were smaller relative to the body, but to the same degree at all body sizes. Selection for large body size led to a change in allometry with a decrease in the allometric coefficient, wing size becoming progressively smaller relative to body as body size increased. When larvae were deprived of food so as to produce adults of a range of small body sizes, all strains retained the same allometric coefficient but showed an increase in the scaling factor. Thus individuals starved as larvae had a smaller adult body size but had proportionally larger wings than fed individuals. We analyzed the developmental processes that could give rise to this pattern of allometries. Differences in the relative growth of body and wing disks can account for the differences in the allometric coefficients among the three body size strains. The change in wing-body allometry at large body sizes was primarily due to an insufficient time period for growth. The available time period for growth of the wing imaginal disks poses a significant constraint on the proportional growth of wings, and thus on the evolution of large body size.

Larval diapause of Monochamus urussovi and photoperiodic effects on larval development in tree bolts

AbstractTo determine the larval diapause and the effect of photoperiod on development in Monochamus urussovi (Coleoptera: Cerambycidae), larvae were reared on Abies sachalinensis and Picea jezoensis logs and bolts. Larvae stopped developing in the final instar at 25°C and 16L : 8D (16 h light and 8 h dark) whereas an exposure to 5°C in the dark (134 days) following acclimation at 12°C under natural daylength led to adult emergence. When larvae were reared under 8L : 16D or 16L : 8D at 25°C with an intervening period of chilling at 5°C in the dark (112 days), a photoperiod of 8L : 16D induced a shorter time required for adult emergence after being returned to 25°C, and smaller adult body size than 16L : 8D.

Survivorship rates of adult Anolis mariarum (Squamata: Polychrotidae) in two populations with differing mean and asymptotic body sizes

We compared adult survivorships in two populations of the lizard Anolis mariarum with different mean and asymptotic body sizes to examine one prediction of age-specific mortality theory; that populations that experience higher adult mortality should exhibit earlier maturation and smaller adult body sizes. We used a maximum likelihood approach to evaluate different survivorship models and model-averaging to estimate survivorship and capture probabilities for each site and sex. Relative tail length did not affect survivorship rates of adults in these two populations, but body size was related to survivorship, with the largest individuals at the time of first capture having lower survivorship rates, so body size was included as a covariate in some of the models examined. Analyses revealed that males at both sites had higher survivorships than females, but there were no differences among the sites in survivorship rates or capture probabilities for either sex. The differences in body sizes documented for these sites still could represent life history adaptations to differences among the sites in mortality rates in the egg or juvenile stages of the life cycle, or may represent a case of phenotypic plasticity to differing environmental conditions, but they appear not to be related to differences in adult survivorships. The estimates of annual survivorships (11.7% to 21.2%) were high for a small, mainland Anolis, and this is the first report of survivorships of male anoles exceeding those of females.

Does harvest select for maladaptation in an increasingly variable world?

For several plant and animal species, Darimont et al. (1) present compelling evidence that rates of phenotypic change driven by human harvest far outpace those driven by other selective forces. From a life-history perspective, commercial and trophy harvesting also resulted in smaller size/earlier age at first reproduction, smaller adult body size (sometimes associated with shorter lifespan), and increased reproductive investment. Selection for these traits can mold life histories with faster reproductive tempos that, not surprisingly, can increase maximum yield (in numbers of individuals) under the assumption of a constant environment. In a world in which climatic conditions are predicted to become increasingly variable (2), however, commercial and trophy harvesting practices may be culling the very traits that would be most adaptive in the future.

Description and Phylogenetic Relationships of Two New Species of Miniature Arthroleptis (Anura: Arthroleptidae) from the Eastern Arc Mountains of Tanzania

I describe two new species of squeaker frog (Arthroleptidae: Arthroleptis) from the Eastern Arc Mountains of Tanzania. These new species are distinguished from other miniature Arthroleptis in the Eastern Arc Mountains by the combination of very small adult body size (< 15 mm snout–vent length), a pronounced dark inguinal spot, and color patterns unique to each species. The new species could be the smallest frog species known from East Africa and among the smallest species of Arthroleptis. An estimate of phylogeny on the basis of mitochondrial DNA sequences reveals that these morphologically cryptic species are sister taxa that compose a basal lineage within a clade of miniature Arthroleptis. Their description hints at a diverse cryptic amphibian fauna in the Eastern Arc Mountains that awaits additional discovery through molecular methods.

Climate change and micro-geographic variation in laying date

Climate change has been shown to affect the timing of reproduction, with earlier reproduction being associated with an increase in temperature. Changes in the timing of reproduction arise from changes in food availability as well as other factors, and differences in the timing of reproduction among sites may cause sites with early reproduction to contribute disproportionately to local recruitment. In this study, spatial variation in the laying date of barn swallows Hirundo rustica at 39 sites in a 45-km2 study area during the period 1971-2004 was used to investigate micro-geographic patterns in the timing of breeding. I found that individuals breeding at sites with early reproduction had a disproportionately large reproductive success. Early sites were characterized by early plant phenology, as determined by the date of leafing of the broad-leaved elm Ulmus glabra and date of flowering of the snowdrop Galanthus nivalis during a single year. Such early sites showed greater advancement in laying date between 1971 and 2004 than the average site. Early sites were also generally occupied during more years by a larger number of breeders than were late sites. Breeders at early sites produced more fledglings, and breeders at such sites were characterized by a smaller adult body size and larger secondary sexual characters than individuals at the average site. These observations are consistent with the hypothesis that temporal changes in the timing of reproduction occur as a consequence of differential recruitment at phenologically early sites that produce disproportionately large numbers of offspring.

Independent evolutionary origins of landlocked alewife populations and rapid parallel evolution of phenotypic traits

Alewife, Alosa pseudoharengus, populations occur in two discrete life-history variants, an anadromous form and a landlocked (freshwater resident) form. Landlocked populations display a consistent pattern of life-history divergence from anadromous populations, including earlier age at maturity, smaller adult body size, and reduced fecundity. In Connecticut (USA), dams constructed on coastal streams separate anadromous spawning runs from lake-resident landlocked populations. Here, we used sequence data from the mtDNA control region and allele frequency data from five microsatellite loci to ask whether coastal Connecticut landlocked alewife populations are independently evolved from anadromous populations or whether they share a common freshwater ancestor. We then used microsatellite data to estimate the timing of the divergence between anadromous and landlocked populations. Finally, we examined anadromous and landlocked populations for divergence in foraging morphology and used divergence time estimates to calculate the rate of evolution for foraging traits. Our results indicate that landlocked populations have evolved multiple times independently. Tests of population divergence and estimates of gene flow show that landlocked populations are genetically isolated, whereas anadromous populations exchange genes. These results support a 'phylogenetic raceme' model of landlocked alewife divergence, with anadromous populations forming an ancestral core from which landlocked populations independently diverged. Divergence time estimates suggest that landlocked populations diverged from a common anadromous ancestor no longer than 5000 years ago and perhaps as recently as 300 years ago, depending on the microsatellite mutation rate assumed. Examination of foraging traits reveals landlocked populations to have significantly narrower gapes and smaller gill raker spacings than anadromous populations, suggesting that they are adapted to foraging on smaller prey items. Estimates of evolutionary rates (in haldanes) indicate rapid evolution of foraging traits, possibly in response to changes in available resources.

Rapid population divergence in thermal reaction norms for an invading species: breaking the temperature–size rule

The temperature-size rule is a common pattern of phenotypic plasticity in which higher temperature during development results in a smaller adult body size (i.e. a thermal reaction norm with negative slope). Examples and exceptions to the rule are known in multiple groups of organisms, but rapid population differentiation in the temperature-size rule has not been explored. Here we examine the genetic and parental contributions to population differentiation in thermal reaction norms for size, development time and survival in the Cabbage White Butterfly Pieris rapae, for two geographical populations that have likely diverged within the past 150 years. We used split-sibship experiments with two temperature treatments (warm and cool) for P. rapae from Chapel Hill, NC, and from Seattle, WA. Mixed-effect model analyses demonstrate significant genetic differences between NC and WA populations for adult size and for thermal reaction norms for size. Mean adult mass was 12-24% greater in NC than in WA populations for both temperature treatments; mean size was unaffected or decreased with temperature (the temperature-size rule) for the WA population, but size increased with temperature for the NC population. Our study shows that the temperature-size rule and related thermal reaction norms can evolve rapidly within species in natural field conditions. Rapid evolutionary divergence argues against the existence of a simple, general mechanistic constraint as the underlying cause of the temperature-size rule.

Hypoxia and seasonal temperature: Short-term effects and long-term implications for Acartia tonsa dana

Far more attention has been given to the short-term lethal impacts of reduced dissolved oxygen on commercially important fish and crabs than to the long-term sublethal impacts on these same species, or on lower trophic levels. This study demonstrates that chronic, sublethal effects of hypoxia on the copepod Acartia tonsa, a critical component of many pelagic coastal food webs, can lead to significant decreases in population growth. The results of laboratory experiments conducted at 15 °C (winter) and 25 °C (summer), under conditions of normoxia (Controls), sublethal hypoxia (1.5 ml l − 1) and lethal hypoxia (0.7 ml l − 1) show that egg production female − 1 day − 1 was significantly lower at 0.7 ml l − 1 compared to Controls at both temperatures, while egg production female − 1 day − 1 was significantly lower at 1.5 ml l − 1 compared to controls in both summer experiments and in one of the two winter experiments. Survival was significantly decreased in the 0.7 ml l − 1 treatment compared to Controls and the 1.5 ml l − 1 treatment. Copepods developed more slowly and matured at smaller adult body sizes at both temperatures under both lethal and sublethal hypoxia compared to normoxia. Under summer temperatures, egg production was reduced by hypoxia exposure on two counts: (1) exposure to hypoxia during development resulted in smaller adults, which translated into lower egg production, and (2) egg production was still significantly lower in hypoxia treatments compared to Controls even when differences in body size were taken into account. While copepods collected in winter and exposed to winter temperatures and hypoxia also matured at smaller body sizes than copepods exposed to normoxia, egg production in winter was almost entirely attributable to this reduction in body size. These results suggest that coastal hypoxia may have a significantly greater impact in the summer months, when copepod populations are most abundant and growing at their most rapid rate of the year. With the anticipated increases in global temperatures, hypoxia may have even greater impacts on pelagic food webs.

Interpopulation variation in growth and life-history traits of the introduced sunfish, pumpkinseed Lepomis gibbosus, in southern England

Summary We examined attributes of growth and reproduction in 19 populations of pumpkinseed (Lepomis gibbosus) introduced into southern England in order to: (i) assess variability of these traits in a northern European climate; (ii) assess inter-relationships among these variables; and (iii) compare these attributes with populations from other parts of Europe where pumpkinseeds have been introduced. Growth rates varied considerably among populations, but juvenile growth rates and adult body sizes were generally among the lowest in Europe. Mean age at maturity ranged from 2.0 to 3.9, and was strongly predicted by the juvenile growth rate (earlier maturity with faster juvenile growth). Other population parameters that also displayed significant negative associations with mean age at maturity were gonadosomatic index, body condition, and adult body size (total length, TL at age 5). Mean TL at maturity and the adult growth increment showed no significant associations with any of the other growth or life-history variables. Pumpkinseed populations in England matured significantly later than those introduced into warmer, more southerly areas of the continental Europe. All of these data suggest that a combination of cool summer temperatures and resource limitation is the cause of slow growth, small adult body size and delayed maturity relative to introduced populations on the European mainland.

Fetal programming: Adaptive life-history tactics or making the best of a bad start?

Fetal programming is an ontogenetic phenomenon of increasing interest to human biologists. Because the downstream consequences of fetal programming have clear impacts on specific life-history traits (e.g., age at first reproduction and the general age-pattern of reproductive investments), a number of authors have raised the question of the adaptive significance of fetal programming. In this paper, I review in some detail several classical models in life-history theory and discuss their relative merits and weaknesses for human biology. I suggest that an adequate model of human life-history evolution must account for the highly structured nature of the human life cycle, with its late age at first reproduction, large degree of iteroparity, highly overlapping generations, and extensive, post-weaning parental investment. I further suggest that an understanding of stochastic demography is essential for answering the question of the adaptive significance of fetal programming, and specifically the finding of low birth weight on smaller adult body size and earlier age at first reproduction. Using a stage-structured stochastic population model, I show that the downstream consequences of early deprivation may be "making the best of a bad start" rather than an adaptation per se. When a high-investment strategy entails survival costs, the alternate strategy of early reproduction with relatively low investment may have higher fitness than trying to play the high-investment strategy and failing.

Temperature dependence of fitness components in geographical populations of Drosophila melanogaster: changing the association between size and fitness

Contrary to the conventional wisdom ‘bigger is better’, evolution at high temperature invariably leads to small individuals in Drosophila melanogaster. Natural selection is known to be responsible, meaning that genotypes that are small because of adaptation to high temperature must have some temperature dependent fitness advantage. In this study we consider both preadult and adult fitness components, and show that large adults from a cold adapted population significantly outperform small adults from a warm adapted population only when tested at low temperature and low larval density. In all other conditions ‘bigger is not necessarily better’, meaning that environmental influences are capable of altering the association between size and fitness. Yet, ‘smaller wasn’t better either’ under any condition, when considering the overall measure of fitness. Examination of individual fitness components revealed population by temperature interaction in preadult survival; this interaction is potentially capable of explaining the temperature specific advantage of small adult body size. At high temperature, the warm adapted population exhibits superior preadult survival while producing small adults. Geographical variation in adult body size seems to be the result of selection on larval growth and competitive strategies, resulting in alterations in the association between fitness components. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 717–725.

The effect of past condition on a multicomponent sexual signal.

Sexual-selection theory predicts that multiple signals may reveal male condition at different stages of life, thus allowing females to make a more reliable assessment of male quality. While the effect of current condition on signal design is well established, few studies have experimentally investigated the effects of past condition. We therefore manipulated the nutritional condition of male nymph field crickets Gryllus campestris and assessed the enduring effects on multiple components of the adult calling song. Food-restricted males had longer nymphal development times and smaller adult body sizes than nymphs with ad libitum food access. Nymphal feeding conditions specifically affected the allometric relationship between body size and harp size, as food-restricted males developed comparatively small harps, leading to a calling song of higher carrier frequency than that produced by similar-sized control males. Other calling-song components, notably chirp rate and chirp intensity, were not affected by the nymphal food treatment, exposing carrier frequency as the key component indicating past condition. In a previous study we established chirp rate as the sole indicator of current condition. The combined results represent experimental evidence of a multicomponent sexual signal that provides distinct information on male condition during different stages of life.

A revision of the Paraulopus oblongus group (Aulopiformes: Paraulopidae) with description of a new species

Paraulopus brevirostris, P. filamentosus, P. japonicus, P. legandi, P. maculatus, and P. oblongus are redescribed. All species are included in the Paraulopus oblongus group, characterized by having no supraocular ridge, 2.5–3.5 scales above the lateral line, and small adult body size (70–150 mm SL). In addition, P. atripes, from the Indian Ocean, is described as a new species of the P. oblongus group, being defined by the following combination of characters: small antrorse dentary process on chin; eye directed laterally; pelvic fin black in males; caudal fin white; 32–34 gill rakers; 45–46 pored lateral-line scales; and 3.5 scales below lateral line. Paraulopus albimaculatus is regarded as a junior synonym of P. brevirostris, based on examination of type specimens. A key to species in the P. oblongus group is included.

Natural selection for grazer resistance to toxic cyanobacteria: evolution of phenotypic plasticity?

We studied the selection response of the freshwater grazing zooplankter, Daphnia galeata, to increased abundance of cyanobacteria in its environment. Cyanobacteria are a poor-quality and often toxic food. Distinct genotypes of D. galeata were hatched from diapausing eggs extracted from three time horizons in the sediments of Lake Constance, Europe, covering the period 1962 to 1997, a time of change in both the prevalence of planktonic cyanobacteria and levels of phosphorus pollution. We assessed whether the grazers evolved to become more resistant to dietary cyanobacteria by exposing genetically distinct clones to two diets, one composed only of the nutritious green alga, Scenedesmus obliquus (good food), and the other a mixture of S. obliquus and the toxic cyanobacterium Microcvstis aeruginosa (poor food). Genotype performance was measured as the specific rate of weight gain from neonate to maturity (gj). We evaluated evolutionary change in the Daphnia population using an analysis of reaction norms based on relative (log-transformed) changes in gj. Log(gj) is a measure of the proportional effect of dietary cyanobacteria on other fitness components of the Daphnia phenotype. For comparison, we also analyze absolute (i.e., nontransformed) changes in gj and discuss the interpretations of the two approaches. Statistical results using a general linear model demonstrate a significant effect of genotype (showing differences in gj among genotypes), a significant genotype x food-type interaction (showing differences in phenotypic plasticity among genotypes), and, in the case of log-transformed data, a significant sediment-genotype-age x food-type interaction. The latter shows that phenotypic plasticity evolved over the period studied. Two constraints act on response to selection in the D. galeata-Lake Constance system. First, gj on a diet containing poor food is highly correlated with gj on a diet of good food, thus evolving resistance also meant evolving an increase in gj on both diets. Second, because genotypes with a high gj also grow to a large adult body size, which in turn increases Daphnia vulnerability to fish predation, we suggest that selection only acted to favor genotypes possessing a high potential gj after cyanobacteria became prevalent. The presence of cyanobacteria depressed realized gj and led to animals of small adult body size even if their genotypes had the potential for high gj and large size. With realized gj reduced, genotypes with an inherently high value could be selected even in the presence of predatory fish. The joint action of selection by dietary cyanobacteria and vulnerability to fish predation provides an explanation for the observed evolution of resistance to poor food through reduced phenotypic plasticity.

Cephalopod consumption by trawl caught fish in Scottish and English Channel waters

Although there have been numerous studies on diets of fish from European waters, they are often of limited use for estimating the impact of fish predation on cephalopod resources. This is due to either the origin of the fish sampled, e.g. from areas with a relatively low cephalopod abundance, or to the method of prey identification: cephalopods are rarely identified to species. This paper presents a summary of recent sampling of the cephalopod species appearing in the diet of commercially important fish species in UK waters. Samples of demersal fish from around Scotland and from the English Channel were collected opportunistically during research trawl surveys and from commercial trawlers in different seasons. For all fish species examined, incidence of cephalopods in the stomachs was low, with fish and crustacean prey dominating stomach contents. Most of the cephalopods found were species with little or no commercial value and small adult body size, such as the small loliginid squid Alloteuthis spp. and bobtail squid (Sepiolidae). Only during December 1998 were some commercially valuable squid ( Loligo spp.) observed in fish stomachs, although in relatively small quantities and comprising mainly of pre-recruit individuals. It is reasonable to suggest that most commercially fished teleosts have little direct impact on post-recruits of commercially important cephalopod stocks, although their impact on recruitment may be significant. The stomach contents data are used to provide a rough estimate of cephalopod consumption by cod populations in the study area and the use of such data to estimate natural mortality is discussed.

NATURAL SELECTION FOR GRAZER RESISTANCE TO TOXIC CYANOBACTERIA: EVOLUTION OF PHENOTYPIC PLASTICITY?

We studied the selection response of the freshwater grazing zooplankter, Daphnia galeata, to increased abundance of cyanobacteria in its environment. Cyanobacteria are a poor-quality and often toxic food. Distinct genotypes of D. galeata were hatched from diapausing eggs extracted from three time horizons in the sediments of Lake Constance, Europe, covering the period 1962 to 1997, a time of change in both the prevalence of planktonic cyanobacteria and levels of phosphorus pollution. We assessed whether the grazers evolved to become more resistant to dietary cyanobacteria by exposing genetically distinct clones to two diets, one composed only of the nutritious green alga, Scenedesmus obliquus (good food), and the other a mixture of S. obliquus and the toxic cyanobacterium Microcystis aeruginosa (poor food). Genotype performance was measured as the specific rate of weight gain from neonate to maturity (gj).We evaluated evolutionary change in the Daphnia population using an analysis of reaction norms based on relative (log-transformed) changes in gj. Log(gj) is a measure of the proportional effect of dietary cyanobacteria on other fitness components of the Daphnia phenotype. For comparison, we also analyze absolute (i.e., nontransformed) changes in gj and discuss the interpretations of the two approaches. Statistical results using a general linear model demonstrate a significant effect of genotype (showing differences in gj among genotypes), a significant genotype × food-type interaction (showing differences in phenotypic plasticity among genotypes), and, in the case of log-transformed data, a significant sediment-genotype-age × food-type interaction. The latter shows that phenotypic plasticity evolved over the period studied.Two constraints act on response to selection in the D. galeata–Lake Constance system. First, gj on a diet containing poor food is highly correlated with gj on a diet of good food, thus evolving resistance also meant evolving an increase in gj on both diets. Second, because genotypes with a high gj also grow to a large adult body size, which in turn increases Daphnia vulnerability to fish predation, we suggest that selection only acted to favor genotypes possessing a high potential gj after cyanobacteria became prevalent. The presence of cyanobacteria depressed realized gj and led to animals of small adult body size even if their genotypes had the potential for high gj and large size. With realized gj reduced, genotypes with an inherently high value could be selected even in the presence of predatory fish. The joint action of selection by dietary cyanobacteria and vulnerability to fish predation provides an explanation for the observed evolution of resistance to poor food through reduced phenotypic plasticity.Corresponding Editor: A. Caballero

Diet, Body Size, and Landscape Use among Holocene People in the Southern Cape, South Africa

Diet, Body Size, and Landscape Use among Holocene People in the Southern Cape, South Africa

Nomenclatural History and Rediscovery of Rhacophorus lateralis Boulenger, 1883 (Amphibia: Rhacophoridae).

The poorly known south-west Indian rhacophorid, Rhacophorus lateralis Boulenger, 1883, known from a unique holotype in the BMNH, is redescribed based on two adult females, from South Coorg, Karnataka State, south-western India. The species is listed in recent lists as valid, despite an attempted synonymy with Rhacophorus malabaricus Jerdon, 1870, by Wolf (1936). The species is diagnosed by the following suite of characters: skin of forehead free; dorsum dark brown with a pair of yellow lines that run from the region around the nostrils, over the eyelids, along the sides of the body, terminating in the inguinal region; small adult body size (SVL to 32.8 mm). The species is illustrated for the first time.