Segregation in spawning and early life history among polymorphic Arctic charr, Salvelinus alpinus, in Thingvallavatn, Iceland

The relationships among time of spawning, incubation temperature, timing of first feeding and early growth were examined in four sympatric morphs of Arctic charr in Thingvallavatn, Iceland. Large benthivorous charr spawn in July-August at sites with cold ground-water flow. Planktivorous and piscivorous charr spawn in September-November and are not confined to ground-water sites. The spawning of small benthivorous charr overlaps with that of other morphs. Progeny of large benthivorous charr start feeding 2-3 months earlier than the progeny of autumn spawners. This results in differential size distribution and growth rates of young in the spring.     

Genetic basis of life-history variation of dwarf and normal Arctic charr,Salvelinus alpinus (L.), in Stora Rösjön,central Sweden*

The genetic basis of life-history variation of dwarf and normal Arctic charr forms in Stora Rosjon was studied by means of two separate rearing experiments on the progeny of artificially bred wild fish. The concept of differential life-strategies gained support, since the progeny of the normal charr form had a higher growth rate and became sexually mature at a larger size and later age than the progeny of the dwarf charr form. The hybrid progeny (dwarf × normal parental charr) was recognized as intermediate in growth as well as in size and age at maturity, compared to the normal and dwarf progeny. However, a dominance effect of the growth and maturation pattern of the dwarf charr was revealed, since hybrid progeny were more similar to dwarf than to normal progeny. The rearing experiments also support the view that the dwarf and normal charr forms are inherited characteristics of one population, since any individual progeny cannot be assigned to a certain form according to its size and age at maturity.     

The functional significance of inherited differences in feeding morphology in a sympatric polymorphic population of Arctic charr

Artificially fertilised eggs from wild-caught Arctic charr parents of two sympatric morphs (benthivorous and planktivorous) from Loch Rannoch, Scotland were reared in the laboratory under identical conditions. During the subsequent 2 years, aspects of their trophic anatomy and feeding behaviour were compared. As previously described for wild-caught fish, charr derived from the benthivorous morph had an increasingly wider mouth gape for a given body length than those derived from the planktivorous morph. The functional significance of these differences in gape was tested by comparing the maximum size of prey that could be handled by each of the two morphs. In both forms, a larger gape enabled larger food particles to be eaten, but the elevation of the regression of maximum prey size on gape was higher in the benthivorous form, indicating the existence of additional morphological and/or behavioural differences influencing the size of prey consumed. When offered a choice between a typical benthic prey item and a typical pelagic food item, charr of benthivorous origin were more likely to feed on the former, whereas those of planktivorous origin were more likely to feed on the latter. Thus inherited differences in gape place constraints on foraging ability and are associated with inherited differences in dietary preference. We conclude that the functional significance of the foraging specialisations indicate a strong selection pressure for the evolution of the divergence and propose that heterochronic growth is the mechanism resulting in the divergence of tropic anatomy.     

Late ontogeny growth acceleration and size form transformations in Transbaikalian Arctic charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> complex: evidence from fin ray cross section growth layers

In some polymorphic populations of Arctic charr in Transbaikalia, an individual can transform from a smaller to larger size form during their lifetime as a result of accelerated growth that follows a period of slow growth and reproduction as a small size form. Alternating periods of slow and fast growth are reflected in growth layer patterns visible in fin ray cross sections. Stained microtome fin ray cross sections were used to reveal the incidence of transformations from one form to another. Data were collected from 14 northern Transbaikalian lakes containing two or three sympatric Arctic charr forms (‘dwarf’, ‘small’, and ‘large’) exhibiting varying levels of morphological separation. Individuals recruited from the dwarf or small form were found in varying proportions among the small and/or large form in 12 lakes. Small or large form charr that grew without noticeable acceleration to the adult size typical of the form or experienced accelerated growth as juveniles prior to maturation were also observed. There were no transformations between sympatric forms that differed in the length and number of gill rakers and in some other meristic characters. Results indicate that in the region under study, transformations of sympatric Arctic charr size forms are a widespread, but not a ubiquitous, phenomenon. Such transformations reflect the plasticity of the developmental channels of the forms. In the course of intra-lacustrine form divergence and genetic differentiation, the frequency of the observed transformations decreases to zero.     

Racial differences in eggs and juveniles of Windermere charr, Salvelinus alpinus

At least four races of charr occur in Windermere, the largest natural lake in England: north basin and south basin autumn spawners, north basin and south basin spring spawners. This study examines racial differences between eggs and juveniles, and relates juvenile size and survival to egg size. There were no major differences between races for egg incubation times and the percentage of eggs hatching successfully, the latter being high (mean values 76–96%) with a negligible proportion of abnormal alevins (<0.8%). Although there were no significant differences in the lengths of the female parents, both eggs and alevins were significantly larger for the autumn spawners than the spring spawners. Size differences in alevins, especially live weight, were positively related to egg size but not female parent size. Mean percentage survival for juveniles attaining the independent feeding stage was higher for the progeny of autumn spawners (32%) than spring spawners (3%). Racial differences in the egg and alevin stages therefore appear to have a significant effect on subsequent survival, and could be ultimately responsible for the relatively small proportion of spring spawners (only 4–6%) in the Windermere population of charr.     

Aggression and growth as an effect of size composition in groups of Arctic charr

Aggressive interactions were initially higher in groups comprised of equally sized large and intermediate Arctic charr Sahelinus alpinus than in groups of small sized individuals and in groups with a normal size distribution. Size composition had no effect on growth.     

Trophic specialization in Arctic charr Salvelinus alpinus (Pisces; Salmonidae): morphological divergence and ontogenetic niche shifts

Lake Thingvallavatn supports four trophic morphs of Arctic charr, Salvelinus alpinus (L.); two of the morphs are benthic (small and large benthivorous charr) one exploits pelagic waters (planktivorous charr) and the fourth is found in both habitats (piscivorous charr). The morphological variation among these morphs was analysed by use of principal component analysis and canonical discriminant analysis. The benihic morphs have a short lower jaw and long pectoral fins. The benthic fish also have fewer gillrakers than the other morphs. Small and large benthivorous charrs attain sexual maturity from 2 and 6 years of age, and at fork lengths from 7 and 22 cm, respectively. Small benthivorous charr retain their juvenile parr marks as adults, have beige ventral colours, and are frequently melanized under the lower jaw. Planktivorous and piscivorous charr attain sexual maturity from 4 and 6 years of age, from fork lengths of 15 and 23 cm, respectively. This phenotypic polymorphism is associated with habitat utilization and diet of the fish, and has probably arisen within the lake system through diversification and niche specialization. The pelagic morphs apparently stem from a single population, and are possibly diversified through conditional niche shifts which affect ontogeny. Juveniles reaching a body length of 23 cm may change from zooplankton to fish feeding. Asymptotic length increases thereby from 20.5 cm in planktivorous charr to 30.2 cm in piscivorous charr. The benthic morphs appear to represent separate populations, although both feed chiefly on the gastropod Lymnaea peregra. Their co-existence seems to be facilitated by size dependent constraints on habitat use. The small morph (asymptotic length 13.3 cm) exploit the interstitial crevices in the lava block substratum, whereas the large morph (asymptotic length 55.4 cm) live epibenthically.     

Inherited differences in head allometry in polymorphic Arctic charr from Loch Rannoch, Scotland

Planktivorous and benthivorous morphs of wild Arctic charr Salelinus alpinus from Loch Rannoch, Perthshire, Scotland, reared in the laboratory from artificially fertilized eggs under identical conditions, were morphologically distinct (based on a series of head measurements) from an early age. For some morphometric characters, these differences became more marked with increasing body size, reflecting differences in the allometric growth patterns of the two forms. These data show that the observed phenotypic differences between these two trophic variants were, to some extent at least, inherited. Thus it is concluded that the mechanisms regulating expression of phenotype in the sympatric morphs of Loch Rannoch are significantly different from the environmentally determined body size polymorphisms reported from Norway but are closer to the more completely divergent forms from Thingvalavatn, Iceland.     

The Importance of Egg Size and Social Effects for Behaviour of Arctic Charr Juveniles

Early behaviour can determine food intake and growth rate with important consequences for life history and survival in fishes. Egg size is known to affect growth rate of young Arctic charr but its influence on the development of behaviour is poorly documented. It is believed that egg size influence on growth and potentially on the behaviour of young fish decreases over time, minimized by the effects of social factors. Shortly after first feeding, we examined differences in mobility and foraging of Arctic charr in relation to egg size and social environment. The behaviour of juveniles from small and large eggs was compared five times over the course of development and in three different experimental settings: long‐term isolation (isolation before hatching), short‐term isolation vs. group rearing and mixed size group vs. homogeneous size groups. Egg size affected foraging behaviour and mobility of fish: fish coming from large eggs were more mobile and foraged more than fish coming from small eggs. Social environment affected foraging behaviour, mobility and space use: fish in a group were more mobile, foraged more and responded faster to food delivery than isolated fish. The interaction of egg size and social effects was seen primarily in foraging activities but did not affect mobility or space use. Large fish in groups foraged more than the three other groups: large fish in isolation, small fish in groups and small fish in isolation. Agonistic behaviour was rarely observed and there was no significant effect of group composition on agonistic behaviour. We discuss the importance of egg size and social effects at early stages of development with a focus on the evolutionary ecology of Arctic charr.     

Habitat use of arctic charrSalvelinus alpinus in Thingvallavatn,Iceland

Synopsis Habitat use by four morphs of arctic charr,Salvelinus alpinus, was investigated in Thingvallavatn, Iceland, by sampling with pelagic and benthic gill nets. Sampling was done in May/June and August/September. Greatest abundance of fish was recorded in the littoral and epipelagic zone in early autumn. Catches were low in early summer. The four morphs are partly segregated in habitat. Small (SB-) and large benthivorous (LB-) charr have a more restricted spatial distribution than piscivorous (PI-), and especially planktivorous (PL-) charr. Both benthivorous morphs are mainly found in the littoral zone, and occur in largest numbers in stony shallows at depths between 0 and 10 m. PL-charr usually dominates in numbers in all habitats. PI-charr is most abundant in epibenthic habitats, although numbers are always low. All morphs are caught in higher numbers at night than during the day, but the diurnal activity difference is highest among SB-charr. The habitat use by different morphs is as may be expected from their morphology and diets. Within the population of PL-charr, young and small fish are more abundant on the bottom than in the pelagic zone, and there is a surplus of females in the pelagic zone. Along the benthic profile, young, small and immature PL-charr are more abundant in deep than in shallow waters. The results are discussed in relation to food supply, competition and predation. Possible reasons for the occurrence of four arctic charr morphs are also discussed.Contribution from the Thingvallavatn project.     

Alternative mating tactics of arctic charr,Salvelinus alpinus,in Thingvallavatn,Iceland

Synopsis During the spawning season the operational sex ratio of the large benthivorous (LB) arctic charr morph in Thingvallavatn is skewed in favour of males which compete intensely on the spawning site. The skewness is caused by males staying longer on the spawning ground than females. LB males employ two mating tactics. Either they guard the female or they attempt sneak-matings with guarded females. The tactics seem to obey a pure conditional strategy where relative size is the important criterion. Generally, the larger males use the guarding tactic and the smaller the sneaking tactic. Assessment of size is a part of the strategy. Relative size influences male behaviour irrespective of which tactic they employ. Guarding is more successful when courting is considered, both with respect to relative frequency and duration of courting acts. Females are aggressive towards the sneakers especially when they are small. This could be interpreted as female choice in favour of large males, but also as a defence against egg predation. A small dwarflike benthivorous morph (SB) is found in the same area. It is mainly active during night and on rare occasions SB-males are seen sneaking into the nest of LB-females. In such cases they are attacked vigorously by both LB sexes.     

Parallel Evolution,not Always so Parallel: Comparison of Small Benthic Charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis>, from Grímsnes and Thingvallavatn,Iceland

Synopsis Iceland is unique in terms of geologically young freshwater systems and rapid adaptations of fresh water fishes to diverse habitats, e.g. lava with ground water flow. Iceland has six species of freshwater fishes, including Arctic charr, Salvelinus alpinus. Previous research has shown great diversity within this species. Four different morphs of Arctic charr are found in one lake, Thingvallavatn, including a small benthivorous charr. Similar populations of small benthic charr are known from several other Icelandic freshwater locations, including Nautavakir in Grímsnes. Our comparison of the small benthic charr morphs in Thingvallavatn and in Grímsnes showed that they are similar in morphology but distinguishable in several characteristics. Small benthic charr in Grímsnes and Thingvallavatn demonstrate similar adaptations and are an example of parallel evolution. However, subtle morphological differences between them indicate further specialized adaptations at each location.     

Body size affects host defensive behavior and progeny fitness in a parasitoid wasp,Lysiphlebus fabarum

This study tested effects of maternal body size on foraging behavior and progeny development in a thelytokous population of Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae). Small and large wasps were reared from first and second instar hosts [black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae)], respectively, and each was provided with a patch (bean leaf disk) containing either 15 small (second instar) or 15 large (fourth instar) hosts for a 30‐min foraging period. Neither body size nor host size affected time allocation to various behaviors within a patch, but second instar aphids produced significantly more mummies than fourth instars. The preferred attack orientation was from the side of the aphid, suggesting wasps were sensitive to the risk of smearing with cornicle secretions. Few wasps developed in fourth instar hosts, suggesting later host instars were somewhat resistant to parasitism. Second instar hosts, the most suitable stage for L. fabarum development, relied more on defensive behavior, specifically kicking and secreting cornicle droplets. Large wasps were more likely to elicit a double cornicle secretion, indicating that aphids graded their response to the size of the attacker. Larger wasps were also more likely to be smeared with cornicle secretion, suggesting they were more vulnerable than small wasps. Although small wasps had smaller eggs than large wasps, there was no effect of maternal egg size on the size of progeny. However, daughters of small females emerged with larger egg loads than daughters of large mothers, and their eggs tended to be slightly smaller, although not significantly. Regression analysis revealed a positive correlation between maternal egg size and progeny developmental time for small and large wasps, and between maternal egg size and progeny egg load for small wasps. These results confirm maternal effects of body size in an aphid parasitoid, and reveal that vulnerability to host behavioral defenses is also body size dependent.     

Growth, maturation and reproductive investment in Arctic charr

Size and rates of growth in a cohort of 1 + Arctic charr housed in standard conditions were tracked over 12 months (December to December) and mature and immature males and females compared retrospectively. In both sexes, maturing fish were larger than non-maturing ones. In males, this size differential was the result of differences in growth in winter and early spring, but not in the remainder of the study period. In females, size differentials resulted mainly from growth rate differences immediately prior to breeding. In females but not in males, gonadosomatic index was predicted by growth rates in the months leading up to maturation, and among the females that matured, faster growing fish produced more eggs. Lipid reserves in July were correlated negatively with growth during the previous 7 months and, in females only, lipid reserves were significantly lower in maturing fish than in non-maturing fish, indicating that mobilization of lipid energy reserves in maturing fish had commenced by this time. Variation in investment in gonadal tissue, measured as gonadosomatic index, was not explained by variation in July lipid reserves for either males or females. However, July lipid reserves were negatively correlated with egg number, so females investing more in ova exhibited greater depletion of lipid reserves. These results are discussed in the context of the relationship between body condition and the onset of maturation in salmonids, relative investment in reproduction and sexual differences in the cost of reproduction.     

Size at maturity of fluvial white-spotted charr, <Emphasis Type="Italic">Salvelinus leucomaenis</Emphasis>, around the Lake Biwa water system varies with habitat size

Size at maturity of fluvial white-spotted charr, Salvelinus leucomaenis, was studied in small headwater tributaries of nine rivers around the Lake Biwa water system, Japan. Threshold size at maturity in both sexes showed significant positive relationships with water discharge, indicating that smaller threshold sizes at maturity of fluvial white-spotted charr occurred in smaller habitats. These results provide a link between size at maturity and habitat size and have important implications for the management of both habitats and white-spotted charr populations.     

Arctic charr in sympatry with burbot: ecological and evolutionary consequences

The trophic niche and parasite infection of Arctic charr (Salvelinus alpinus) were explored in two lakes with sympatric burbot (Lota lota) and two lakes without burbot in subarctic Norway. The CPUE of burbot and charr were similar in one lake, but burbot had a low population density in the other. Burbot were benthivorous in both lakes. Other co-occurring species like brown trout (Salmo trutta), Atlantic salmon parr (Salmo salar), grayling (Thymallus thymallus) and minnow (Phoxinus phoxinus) were also benthivores. At high densities, benthivorous burbot forced the whole Arctic charr population to utilise mainly the limnetic trophic niche. In contrast, at low burbot density or without burbot present, Arctic charr were primarily benthivorous in the littoral zone. Thus, a clear interactive segregation in diet was observed between Arctic charr and burbot at high burbot densities. There was also a high predation pressure from burbot on young Arctic charr along the benthic zones. The extensive use of zooplankton as prey caused a high parasite infection pressure of copepod transmitted Diphyllobothrium spp. larvae, with the potential for high negative impact on the Arctic charr population. As the benthivore trophic niche was occupied by burbot, the ecological opportunities for polymorphism with benthivorous ecotypes or morphs of Arctic charr were probably prevented. Therefore, the sympatry with burbot seems to have large ecological and evolutionary consequences for this Arctic charr population compared with neighbouring lakes where burbot is absent.     

Cryptic maternal effects in Hippodamia convergens vary with maternal age and body size

Maternal effects can mold progeny phenotypes in various ways and may constitute ecological adaptations. By examining the effect of oviposition sequence on progeny produced by different size classes of female ladybird beetles (produced by controlling larval access to food), we show that maternal signals can change through adult life and alter the developmental programs of progeny, ostensibly to synchronize their life histories with predictable resource dynamics, thus maximizing maternal fitness. We also show that female body size, as determined by larval food supply, interacts with female age to influence progeny fitness. When fed ad libitum as adults, small females reared with limited food access laid fewer, smaller eggs than large females reared with ad libitum food access. Maternal body size interacted with oviposition sequence to influence progeny development, but the latter had greater impact. Eggs laid later by medium and large females hatched faster than those laid earlier, larvae fed longer in the fourth instar, their pupation period was shorter, total developmental time was reduced, and adults emerged with greater mass, most notably daughters. Oviposition sequence effects on progeny from small mothers were non‐significant for total developmental time and progeny mass. Only large mothers increased egg size over time and egg mass was not consistently correlated with developmental parameters, indicating that progeny phenotype was impacted by other, more cryptic, maternal signals. Such signals appear costly, as food limitation during development constrained not only fecundity and egg size but also maternal ability to manipulate progeny phenotype. The production of faster‐developing offspring that mature to larger sizes late in the oviposition cycle may be adaptive for exploitation of ephemeral aphid outbreaks with predictable dynamics of prey abundance and competition.     

Ontogenetic variation in fin ray segmentation between latitudinal populations of the medaka, <Emphasis Type="Italic">Oryzias latipes</Emphasis>

Fin rays of ray-finned fishes are composed of multiple bony segments, and each fin ray elongates by adding a new segment to the tip. Therefore, fin ray length is determined by the number of segments and the length of each segment. A comparison of the anal fin rays of a northern and southern wild population of the medaka, Oryzias latipes, revealed that southern fish had more segments per fin ray, resulting in longer anal fins than the northern fish. When fish were reared in a laboratory common environment, segmentation of the fin rays started earlier with respect to body size in the southern fish. In the southern males, moreover, the rate of segment addition accelerated after a certain body size, indicating sexual maturity. These patterns of segment addition during ontogeny were consistent with the patterns of fin ray elongation. Although distal segments tended to be longer, except for the most proximal segment, in both populations, the southern fish had shorter segments than the northern fish at any position on fin rays. These results indicate that the interpopulation variation in fin length is largely due to genetically-based differences in the control of segment addition, and that the length of each segment does not contribute to it. We suspect that fin ray segmentation is regulated by thyroid and sex hormones that differ between populations. We also found that some segments fuse with each other at the base of each fin ray, the functions and mechanisms of which remain unclear.     

Host plant trichomes and the advantage of being big: progeny size variation of the pipevine swallowtail

1. Plants possess numerous traits that confer resistance against insect herbivores, and herbivores, in turn, can evolve traits to ameliorate the effectiveness of these traits. The pipevine swallowtail, Battus philenor, is an extreme specialist on plants in the genus Aristolochia. The only host plant available to the California population of B. philenor is A. californica. Aristolochia californica is distinct from most other B. philenor host plants in that it is pubescent. 2. The progeny of B. philenor are larger in California compared with populations examined in Texas. Size differences persist throughout larval development. 3. Regardless of maternal host plant, population differences in progeny size persist, and crosses between California (large progeny) and Texas (small progeny) B. philenor populations resulted in offspring producing intermediate sized progeny, indicating a heritable component to progeny size variation. 4. California neonate caterpillars more easily overcame the trichomes of A. californica compared with Texas neonates. When trichomes were removed from A. californica, time to feeding establishment was reduced for caterpillars from both populations. Texas caterpillars established feeding sites on A. californica with trichomes removed, in the same time required to establish feeding on their non‐pubescent host plant, A. erecta. 5. This study shows that plant trichomes might impose selection pressure on progeny size.     

Size‐dependent reproductive success of wild zebrafish Danio rerio in the laboratory

Size‐dependent reproductive success of wild zebrafish Danio rerio was studied under controlled conditions in the laboratory to further understand the influence of spawner body size on reproductive output and egg and larval traits. Three different spawner size categories attained by size‐selective harvesting of the F₁‐offspring of wild D. rerio were established and their reproductive performance compared during a 5 day period. As to be expected, large females spawned more frequently and had significantly greater clutch sizes than small females. Contrary to expectations, small females produced larger eggs when measured as egg diameter with similar amounts of yolk compared to eggs spawned by large spawners. Eggs from small fish, however, suffered from higher egg mortality than the eggs of large individuals. Embryos from small‐sized spawners also hatched later than offspring from eggs laid by large females. Larval standard length (L_S)‐at‐hatch did not differ between the size categories, but the offspring of the large fish had significantly larger area‐at‐hatch and greater yolk‐sac volume indicating better condition. Offspring growth rates were generally similar between offspring from all size categories, but they were significantly higher for offspring spawned by small females in terms of L_S between days 60 and 90 post‐fertilization. Despite temporarily higher growth rates among the small fish offspring, the smaller energy reserves at hatching translated into lower condition later in ontogeny. It appeared that the influence of spawner body size on egg and larval traits was relatively pronounced early in development and seemed to remain in terms of condition, but not in growth, after the onset of exogenous feeding. Further studies are needed to explore the mechanisms behind the differences in offspring quality between large‐ and small‐sized spawners by disentangling size‐dependent maternal and paternal effects on reproductive variables in D. rerio.