Population differences in host use by a seed-beetle: local adaptation, phenotypic plasticity and maternal effects

For insects that develop inside discrete hosts, both host size and host quality constrain offspring growth, influencing the evolution of body size and life history traits. Using a two-generation common garden experiment, we quantified the contribution of maternal and rearing hosts to differences in growth and life history traits between populations of the seed-feeding beetle Stator limbatus that use a large-seeded host, Acacia greggii, and a small-seeded host, Pseudosamanea guachapele. Populations differed genetically for all traits when beetles were raised in a common garden. Contrary to expectations from the local adaptation hypothesis, beetles from all populations were larger, developed faster and had higher survivorship when reared on seeds of A. greggii (the larger host), irrespective of their native host. We observed two host plant-mediated maternal effects: offspring matured sooner, regardless of their rearing host, when their mothers were reared on P. guachapele (this was not caused by an effect of rearing host on egg size), and females laid larger eggs on P. guachapele. This is the first study to document plasticity by S. limbatus in response to P. guachapele, suggesting that plasticity is an ancestral trait in S. limbatus that likely plays an important role in diet expansion. Although differences between populations in growth and life history traits are likely adaptations to their host plants, host-associated maternal effects, partly mediated by maternal egg size plasticity, influence growth and life history traits and likely play an important role in the evolution of the breadth of S. limbatus’ diet. More generally, phenotypic plasticity mediates the fitness consequences of using novel hosts, likely facilitating colonization of new hosts, but also buffering herbivores from selection post-colonization. Plasticity in response to novel versus normal hosts varied among our study populations such that disentangling the historical role of plasticity in mediating diet evolution requires the consideration of evolutionary history.     

Phenotypic plasticity in age and size at maturity and its effects on the integrated phenotypic expressions of life history traits of Cardamine flexuosa (Cruciferae)

We analyzed variation in phenotypic plasticity of life history traits between two Cardamine flexuosa populations based on differences in plasticity of age and size at maturity. C. flexuosa (Cruciferae) is a facultative, vernalization-sensitive, long-day annual, and its phenology and the phenotypic expressions of many life history traits are largely controlled by photoperiod and vernalization in natural populations. We used plants from two populations which differed in their responses to chilling and photoperiod treatments. The timing of developmental processes was changed by controlling temperature and photoperiod regimes in growth chambers. Plasticity in size at maturity was analyzed as changes in a growth trajectory using two parameters, age at maturity (Δt) and growth rate (k). Both traits showed plasticity, but differences between the populations were found mostly for Δt. Distinctive differences in size at maturity of individuals in the two populations were mainly due to different amounts of plasticity in Δt. Variations in plasticity of nine other life history traits and their associations to age and size at maturity were also analyzed. Variation for eight of the traits can be described, at least in part, as a function of age and size at maturity for both populations, and most of the variation in the total number of seeds was explained by age and size at maturity. Only age at maturity had any effect on changes in resource allocation. The nine life history traits were integrated through associated character expressions with age and size at maturity. Changes in the association between a trait and age and/or size at maturity were rather conservative compared to changes in the plasticity of a trait between the two populations. Associations with age and size at maturity are mostly explicable in terms of inherent relationships in the developmental processes, and they may limit the ecological range expansion and the adaptive evolution of plasticity in C. flexuosa. The negative correlation between reproductive allocation and age at maturity can be a cost of delaying maturation in C. flexuosa.     

The multiplication of Longidorus elongatus (de Man) on different host plants with reference to virus transmission

In pot tests, Longidorus elongatus (de Man) populations increased rapidly on Fragaria vesca, Stellaria media, Mentha sativa and Lolium perenne and slightly on Brassica rapa (turnip var. White globe) and on Ribes nigrum; but on Rubus idaeus populations declined at the same rate as in fallow soil. Raspberry ringspot (RRV) and tomato black ring (TBRV) viruses were readily transmitted by L. elongatus to S. media, TBRV only was transmitted to L. perenne, and M. sativa became infected with neither virus. RRV was transmitted to R. idaeus var. Mailing Jewel, on which the nematode fed but evidently did not reproduce. In a field experiment L. perenne and Fragaria ananassa (strawberry) vars. Huxley and Redgauntlet were shown to be good hosts for L. elongatus and R. idaeus (raspberry) vars. Lloyd George and Mailing Jewel were confirmed as non-hosts, together with Rheum rhaponticum (rhubarb). Two periods of egg laying were detected each year in the Redgauntlet strawberry plots and these coincided with the growth of young roots during the late spring and autumn. The wide host range of L. elongatus among cultivated plants and weeds, together with its ability to survive long periods without food, precludes the use of crop rotation as a means of control. Chemical soil sterilization is considered the most effective means of control because it can greatly decrease L. elongatus populations and as the nematode multiplies slowly, even on favourable host plants, several years are likely to elapse before populations become large enough to spread viruses effectively from infector plants to susceptible crops.     

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.     

Observations on the distribution patterns,behaviour, diets and reproductive cycles of sand-dwelling clinids (Perciformes: Clinidae) from South Africa

Synopsis Twenty-two samples of sand-dwelling fishes were collected from sublittoral sandy substrata off the Cape Peninsula, South Africa using a novel quantitative technique in which the ichthyocide, rotenone, was introduced beneath 6.25 m² plastic sheets weighted around the perimeter with chain. A total of 94 fish of eight species and four families were recorded. Of these, four species of the family Clinidae made up 92% of the material. Fish densities varied from 0.10–2.96 fish m^–2 (0.17–1.14 g m^–2). Distribution patterns amongst the Clinidae were explained by sediment particle size, with all of the newly discovered species, Cancelloxus longior, inhabiting fine sand (median grain size 0.25–0.50 mm), and all Xenopoclinus leprosus, C. elongatus and Pavoclinus smalei occurring in gravel (> 1 mm). Xenopoclinus kochi was found in all sediment types, but was most abundant in coarse substrata. Diets of all four species were similar, consisting chiefly of amphipods and isopods. However, small differences in prey preference were evident. All four species reached sexual maturity at 25–30 mm standard length, were viviparous, exhibited superembryonation and gave birth to live young of approximately 14 mm. Breeding seasonality occurred in all species except P. smalei.     

Good food versus bad food: the role of sterols and polyunsaturated fatty acids in determining growth and reproduction of <Emphasis Type="Italic">Daphnia magna</Emphasis>

The absence of dietary sterols and polyunsaturated fatty acids (PUFAs) has been shown to affect the performance of the freshwater herbivore Daphnia. Here, we compared somatic growth rates and clutch sizes of Daphnia magna reared on a diet of low food quality (Synechococcus elongatus) and of high food quality (Cryptomonas sp.) and investigated if and to what extent the absence of sterols or PUFAs in the cyanobacterium S. elongatus accounts for the observed differences in food quality. The supplementation of S. elongatus with cell-free lipid extracts (fatty acids, sterols, total lipids) obtained from the flagellate Cryptomonas sp. suggested that the superior food quality of Cryptomonas sp. is predominantly, but not completely, a combined effect of its sterol and PUFA composition. Our laboratory study suggests that somatic growth of D. magna feeding on S. elongatus is primarily constrained by the absence of sterols, whereas egg production is primarily limited by the absence of long chain PUFAs.     

Larval host plant affects fitness consequences of egg size variation in the seed beetle Stator limbatus

Egg size variation often has large effects on the fitness of progeny in insects. However, many studies have been unable to detect an advantage of developing from large eggs, suggesting that egg size variation has implications for offspring performance only under adverse conditions, such as during larval competition, periods of starvation, desiccation, or when larvae feed on low-quality resources. We test this hypothesis by examining the consequences of egg size variation for survivorship and development of a seed-feeding insect, Stator limbatus, on both a low-quality (Cercidium floridum) and a high-quality (Acacia greggii) host plant. Our results are consistent with the hypothesis. S. limbatus larval performance was affected by egg size only when developing on the poor-quality host (C. floridum); larvae from large eggs survived better on C. floridum than those from small eggs, while there was no evidence of an effect of egg size on progeny development time, body weight, or survivorship when larvae developed on A. greggii. These results indicate intense selection for large eggs within C. floridum-associated populations, but not in A. greggii-associated populations, so that egg size is predicted to vary among populations associated with different hosts. Our results also support this hypothesis; females from a C. floridum-associated population (Scottsdale) laid larger eggs than females from an A. greggii-associated population (Black Canyon City).     

Egg size variation between the fluvial-lacustrine and lacustrine types of a landlocked <Emphasis Type="Italic">Rhinogobius</Emphasis> goby in the Lake Biwa water system

The egg size of a landlocked goby, Rhinogobius sp. (the orange form), in the Lake Biwa water system was compared between the fluvial-lacustrine and lacustrine types. The former spawned larger eggs than the latter in the laboratory under the same environmental conditions of water temperature and current. This difference in egg size was also found in the field among six local populations from tributary rivers and lake shore. These results suggested that egg size is largely associated with the life history type that has experienced a different selection regimen, such as the risk of larval starvation, associated with their respective life cycles.     

Photoperiod affects compensating developmental rate across latitudes in the damselfly Lestes sponsa

1. Although there is a great deal of theoretical and empirical data about the life history responses of time constraints in organisms, little is known about the latitude‐compensating mechanism that enables northern populations' developmental rates to compensate for latitude. To investigate the importance of photoperiod on development, offspring of the obligatory univoltine damselfly Lestes sponsa from two populations at different latitudes (53°N and 63°N) were raised in a common laboratory environment at both northern and southern photoperiods that corresponded to the sites of collection. 2. Egg development time was shorter under northern photoperiod regimes for both populations. However, the northern latitude population showed a higher phenotypic plasticity response to photoperiod compared with the southern latitude population, suggesting a genetic difference in egg development time in response to photoperiod. 3. Larvae from both latitudes expressed shorter larval development time and faster growth rates under northern photoperiod regimes. There was no difference in phenotypic plastic response between northern and southern latitude populations with regard to development time. 4. Data on field collected adults showed that adult sizes decreased with an increase in latitude. This adult size difference was a genetically fixed trait, as the same size difference between populations was also found when larvae were reared in the laboratory. 5. The results suggest phenotypic plasticity responses in life history traits to photoperiod, but also genetic differences between north and south latitude populations in response to photoperiod, which indicates the presence of a latitudinal compensating mechanism that is triggered by a photoperiod.     

Hypoxia and life‐history traits in a eurytopic African cichlid

This study quantified variation in key life‐history traits of the widespread African mouth‐brooding cichlid Pseudocrenilabrus multicolor victoriae. Egg size, number, batch reproductive effort, size at maturity and brooding efficiency were compared among field populations across a wide range of dissolved oxygen (DO) concentrations from extreme hypoxia to normoxia. In the laboratory, a similar suite of characters was quantified in F1 of low‐ and high‐DO origin reared under low or high DO. In general, females from low‐DO habitats and females reared under low DO were characterized by a smaller size at maturity and no difference in batch reproductive effort when compared with females from high‐DO habitats or females reared under normoxia. A trade‐off between egg size and number was evident in the field and in the laboratory‐rearing experiment, but the direction of the trade‐off differed. Egg size was negatively correlated with egg number across field populations; females collected from low‐DO sites generally had more, smaller eggs relative to females from high‐DO sites. In the laboratory‐rearing experiment, F1 females of high‐DO origin produced larger, fewer eggs than F1 females of low‐DO origin, lending support to the field results and suggesting a heritable component to these traits. There was also an element of developmental plasticity, F1 females raised under low DO produced larger, fewer eggs compared with F1 females raised under high DO (regardless of population) suggesting that DO may interact with other variables to determine egg size in the field.     

Toxicity ofParthenium hysterophorus extracts toChlorella vulgaris andSynechococcus elongatus

Aqueous extracts from leaf or inflorescence ofParthenium hysterophorus were either algistatic or algicidal toChlorella vulgaris andSynechococcus elongatus. Root extract, however, enhanced the growth of the two algae, but a 2.5% level was algistatic toS. elongatus.     

Sexual differences in growth strategies of the wolf spider Hygrolycosa rubrofasciata

In invertebrates, the size at maturation is considered to be important for adult fitness. In the wolf spider Hygrolycosa rubrofasciata, however, it is only females that clearly benefit of larger size through augmented egg production, while male mating success is determined by display activity not related to size. Thus, we can expect conflicting growth patterns for the sexes. Additionally, populations differ greatly in adult size: individuals from dry habitats are smaller than those from wet habitats. To study the sexual differences in reaction norms of growth, we reared spiderlings from seven populations at two food levels under controlled laboratory conditions and compared size at sexual maturity. The shapes of reaction norms for adult size differed between the sexes. In females, the reaction norms were parallel, but individuals from dry habitats tended to grow larger at the given food levels. In males, there was a significant interaction between food level and population without any consistent differences between populations. Maturation time was a plastic character in both sexes with no genetic differences among populations. However, females on low food level matured later and significantly smaller in size than those on high food level. Males also matured later on low food level, but they were nearly of the same size as males that received more food. Female growth patterns reflected the strong selection for large size at maturity. However, the patterns for males were highly variable, which could be explained by the weak overall selection on male size, which means that any environmental factors can affect male growing patterns. This revised version was published online in November 2006 with corrections to the Cover Date.     

Predation of eggs ofSitona hispidulus [Col.: Curculionidae] in alfalfa

The predation on egg populations ofSitona hispidus (Fabricius) [Col.: Curculionidae] in a stand of alfalfa was evaluated in 1982 and 1983. By using pitfall traps, catches of reproductive adults ofS. hispidulus were correlated with catches of carabid species, and potential egg predators were identified for further evaluation. Results indicated that predators removed 28 % of eggs under field conditions.Amara aenea DeGeer was found to be particularly efficient on eggs ofS. hispidulus under caged-field conditions.      

Reproductive biology of blue sucker in a large Midwestern river

Efforts to protect or rehabilitate depressed blue sucker Cycleptus elongatus populations require an understanding of life‐history characteristics and reproductive biology to provide fisheries managers with the tools required for science‐based management. The objective of this study was to examine the reproductive biology of blue sucker in the Wabash River, Indiana, during March and April 2006. A total of 105 reproductively mature blue sucker (53 males, 52 females) was collected using boat electrofishing to examine size‐at‐maturity, absolute fecundity, gonadosomatic index (GSI), relative fecundity, and estimated egg size. Size‐at‐maturity was estimated at 515 mm total length (TL) for males and 568 mm TL for females. Mean absolute fecundity of females captured during the study was 150 704 eggs per female (range, 26 829–267 471 eggs per female) and was positively related to both TL (r² = 0.66) and wet weight (r² = 0.77). Mean GSI was 6.4% (range, 2–9.3%) for males and 17.2% (range, 4.3–23.4%) for females. Relative fecundity ranged from 15 331 to 65 887 eggs kg^?1 body weight (mean = 46 946 eggs kg^?1 body weight) and was strongly correlated with GSI (r² = 0.87). Mean estimated egg size was 278 eggs g^?1 (range, 229 364 eggs g^?1) and exhibited an inverse relationship to GSI (r² = 0.42). The results of our study provide information on the reproductive biology of blue sucker which can be used to aid in the identification of potential recovery threats for depressed populations.     

Effect of two microalgae concentrations on body size and egg size of the rotifer <Emphasis Type="Italic">Brachionus calyciflorus</Emphasis>

The rotifer, Brachionus calyciflorus, was grown with two algae species (Chlorella sp. and Scenedesmus obliquus) at different concentrations (0.1, 1 and 10 × 10⁶ cells ml⁻¹). The body size (lorica biovolume) of individual rotifer and their egg size were measured when the populations were roughly in the exponential phase of population growth. The body size of the rotifers differed significantly (P < 0.05) among the two algae species used, however this effect was not observed for egg size. The body size of rotifers fed on higher densities of Chlorella sp. (10 × 10⁶ cells ml⁻¹) was significantly larger than for those fed on lower and medium densities (0.1 and 1 × 10⁶ cells ml⁻¹). Body size and egg size of rotifers fed with different amounts of Scenedesmus did not differ significantly. The egg size was significantly larger at higher food level of Chlorella. A significantly positive correlation was observed between the adult rotifer body size and their egg size.     

Characterization of gastrointestinal chitinase in the lizard Sceloporus undulatus garmani (Reptilia: Phrynosomatidae)

Most studies on chitinase activity in lizards have been concerned with Palaearctic (European) and Laurasian (Middle Eastern and Asian) taxa. Several genera of Old World lizards, Anguis, Uromastix, Chamaeleo and Lacerta, have been shown to possess chitinolytic activity. To date, only one New World lizard, Anolis carolinensis, has been reported to exhibit chitinolytic activity. In the present study, chitinase activity was characterized in a second New World taxon, Sceloporus undulatus garmani, a New World, phrynosomatid lizard. Chitinolytic activity was measured by incubating tissue extracts with a radioactive chitin substrate, acetyl-[H³]chitin and determining acid soluble radioactivity as an estimate for chitin hydrolysis. Chitinolytic activity was present in stomach, small intestine and pancreas extracts, with the stomach and pancreas having the highest specific activities. Chitinolytic activity was higher at pH 4.5 than at pH 7.5. The stomach chitinase is immunologically similar to the gastric chitinase previously described for rainbow trout. Western blot analysis showed anti-chitinase cross-reactivity in the extracts of the stomach, but no cross-reactivity in the pancreatic or intestinal extracts, suggesting different isoforms of chitinase. There was no detected lysozyme activity (less than 0.01 mg/ml lysozyme) present in the extracts of the stomach, small intestine and pancreas. The localization of chitinolytic activity in S. u. garmani is in agreement with earlier reptilian reports on the distribution of chitinase.     

Female‐biased dimorphism in size and age at maturity is reduced at higher latitudes in lake whitefish Coregonus clupeaformis

Female‐biased sexual dimorphism in size at maturity is a common pattern observed in freshwater fishes with indeterminate growth, yet can vary in magnitude among populations for reasons that are not well understood. According to sex‐specific optimization models, female‐biased sexual size dimorphism can evolve due to sexual selection favouring earlier maturation by males, even when sexes are otherwise similar in their growth and mortality regimes. The magnitude of sexual size dimorphism is expected to depend on mortality rate. When mortality rates are low, both males and females are expected to mature at older ages and larger sizes, with size determined by the von Bertalanffy growth equation. The difference between size at maturity in males and females becomes reduced when maturing at older ages, closer to asymptotic size. This phenomenon is called von Bertalanffy buffering. The predicted relationship between the magnitude of female‐biased sexual dimorphism in age and size at maturity and mortality rate was tested in a comparative analysis of lake whitefish Coregonus clupeaformis from 26 populations across a broad latitudinal range in North America. Most C. clupeaformis populations displayed female‐biased sexual dimorphism in size and age at 50% maturity. As predicted, female‐biased sexual size dimorphism was less extreme among lower mortality, high‐latitude populations.     

Ontogenic colour change,survival, and mating in the damselfly Agriocnemis pygmaea Rambur (Insecta: Odonata)

1. Damselflies often show intra-specific colour variation, which may represent genetic polymorphism or age-related (ontogenic) colour changes. 2. Such variation has distinct implications for the species' ecology and evolution. Colour variation in females of the damselfly Agriocnemis pygmaea was studied, which range from blue male-like individuals (andromorphs) to those with a distinct red colour (heteromorphs). From preliminary observations, it was hypothesised that this species exhibits ontogenic colour change from heteromorph to andromorph coloration. 3. Mark–recapture experiments and egg counts of dissected females suggested that immature females are heteromorphic and gradually begin to resemble males as they attain sexual maturity. 4. Reflectance spectra of field-caught individuals indicated that, although males are indistinguishable from andromorphs, they could be easily differentiated from heteromorphs. 5. Finally, field observations and mate choice experiments showed that males rarely attempt to mate with heteromorphic females and prefer andromorphs. Together, this study's results suggest that the observed colour variation in A. pygmaea females is ontogenic and is associated with sexual maturity.     

Evolutionary divergence in life history traits among populations of the Lake Malawi cichlid fish Astatotilapia calliptera

During the early stages of adaptive radiation, populations diverge in life history traits such as egg size and growth rates, in addition to eco‐morphological and behavioral characteristics. However, there are few studies of life history divergence within ongoing adaptive radiations. Here, we studied Astatotilapia calliptera, a maternal mouthbrooding cichlid fish within the Lake Malawi haplochromine radiation. This species occupies a rich diversity of habitats, including the main body of Lake Malawi, as well as peripheral rivers and shallow lakes. We used common garden experiments to test for life history divergence among populations, focussing on clutch size, duration of incubation, egg mass, offspring size, and growth rates. In a first experiment, we found significant differences among populations in average clutch size and egg mass, and larger clutches were associated with smaller eggs. In a second experiment, we found significant differences among populations in brood size, duration of incubation, juvenile length when released, and growth rates. Larger broods were associated with smaller juveniles when released and shorter incubation times. Although juvenile growth rates differed between populations, these were not strongly related to initial size on release. Overall, differences in life history characters among populations were not predicted by major habitat classifications (Lake Malawi or peripheral habitats) or population genetic divergence (microsatellite‐based F_ST). We suggest that the observed patterns are consistent with local selective forces driving the observed patterns of trait divergence. The results provide strong evidence of evolutionary divergence and covariance of life history traits among populations within a radiating cichlid species, highlighting opportunities for further work to identify the processes driving the observed divergence.     

The origin and development of individual size variation in early pelagic stages of fish

Size variation among individuals born at the same time in a common environment (within cohorts) is a common phenomenon in natural populations. Still, the mechanisms behind the development of such variation and its consequences for population processes are far from clear. We experimentally investigated the development of early within-cohort size variation in larval perch (Perca fluviatilis). Specifically we tested the influence of initial variation, resulting from variation in egg strand size, and intraspecific density for the development of size variation. Variation in egg strand size translated into variation in initial larval size and time of hatching, which, in turn, had effects on growth and development. Perch from the smallest egg strands performed on average equally well independent of density, whereas larvae originating from larger egg strands performed less well under high densities. We related this difference in density dependence to size asymmetries in competitive abilities leading to higher growth rates of groups consisting of initially small individuals under high resource limitation. In contrast, within a single group of larvae, smaller individuals grew substantially slower under high densities whereas large individuals performed equally well independent of density. As a result, size variation among individuals within groups (i.e. originating from the same clutch) increased under high densities. This result may be explained by social interactions or differential timing of diet shifts and a depressed resource base for the initially smaller individuals. It is concluded that to fully appreciate the effects of density-dependent processes on individual size variation and size-dependent growth, consumer feedbacks on resources need to be considered.