Cyclomorphosis in Daphnia galeata mendotae Birge and D. retrocurva Forbes as a predator-induced response

SUMMARY. 1. Populations of Daphnia galeata mendotae and D. retrocurva were exposed for one generation to predators enclosed in mesh bags. The exposed Daphnia showed adaptive morphological changes that mimic cyclomorphosis in nature. The invertebrate predators Chaoborus and Notonecta induced longer helmets, while the fish Lepomis induced shorter bodies but had no effect on helmet length. Longer tail spines (relative to body length) were induced by Notonecta and Lepomis.
2. The responses of D. retrocurva were influenced by algae concentration, with the more extreme responses occurring at a higher food concentration and higher lipid index.     

Extreme helmet formation in Daphnia cucullata induced by small-scale turbulence

Although Daphnia cucullata is used as a textbook example forcyclomorphosis, distinct helmet development, as shown in thefield, has not been demonstrated in the laboratory until now.We show for the first time that small-scale turbulence is ableto induce the maximum response of morphological plasticity inD. cucullata. Helmet elongation reached magnitudes as extremeas observed in the field. Different modes of generating small-scaleturbulence caused different levels of helmet formation. Small-scaleturbulence alone may not be responsible for cyclomorphosis innature because field data show that two nearby lakes with similarmorphometry differ in cyclomorphosis patterns, while laboratoryexperiments show that there is no difference in the abilityto form helmets in the clones of each lake. Although helmetformation in D. cucullata is inducible with predator kairomones,helmet elongation is not as strong as that induced by turbulence.We discuss the possible role of helmets under turbulent conditions.     

Clutch size variation and morphology in a cyclomorphic Bosmina population

Clutch size, cyclomorphosis and allometric growth were analysedin a population of the humpbacked Bosmina (E.) coregoni gibbera.This species shows cyclomorphosis in antennule length and bodyheight, traits that may reduce predation risk from Leptodorakindtii. Individuals with long antennule and extreme body heightmay pay a cost in terms of decreased reproductive capacity.On the other hand, increasing the body height may be a way toincrease the brood chamber volume during periods of good foodconditions. We tested these hypotheses by comparing the seasonalvariation in clutch size with the seasonal variation in morphology.Antennule length and body height increased from mid-May to Augustand showed usually positive allometry at times with high populationdensities of the predatory cladoceran Leptodora kindtii. Clutchsize decreased from spring to late summer contrary to the hypothesisthat cyclomorphosis in height is caused by a seasonal variationin reproductive demand. However, within-dates antennule lengthwas negatively related and body height was positively relatedto clutch size. We conclude that the long antennule may imposea cost that reduces the reproductive capacity. The hypothesisthat carapace cyclomorphosis is driven by seasonally varyingclutch sizes was rejected.     

Genotypic succession in the cyclomorphosis of Bosmina longirostris (Cladocera)

SUMMARY. The pattern of cyclomorphosis of Bosmina longirostris was followed over a 21-month period in Lake Travis, Texas. Appreciable variation was observed in body size and in the lengths of antennules and mucrones. Electrophoresis of clonal isolates collected over the period of study showed a seasonal cycle in esterase phenotypes. A definite association was demonstrated between the esterase phenotypes and morphologies of these clonal isolates. Individuals of one esterase phenotype were usually large and long-featured, whereas individuals of the other esterase phenotypes were smaller, with short features. The association of morphology and esterase phenotypes from laboratory experiments corresponded to the association observed in field samples. These results support the hypothesis that the cyclomorphosis of Lake Travis Bosmina involves a succession of clones as well as phenotypic plasticity.     

Induction and costs of tail spine elongation in Daphnia hyalina×galeata: reduction of susceptibility to copepod predation

1. Seasonal field data showed a positive correlation between the tail spine length of Daphnia hyalina × galeata and the density of the copepod Acanthocyclops robustus . Laboratory experiments were designed to assess the mechanisms underlying the induction of tail spine elongation and to test whether this morphology reduced predation on Daphnia juveniles.
2. Both the elongated tail spine morph and control spine morph produced progeny with elongated tail spine when exposed to copepods. Instar increments, calculated from individual body length, showed that individual growth decreased in the presence of copepods. This decrease in individual growth was less pronounced in spined morphs relative to control morphs.
3. Copepods exhibited significantly higher feeding rates on control morph juveniles compared to elongated tail spine morph juveniles.
4. All Daphnia matured at instar 5 even though the control morph neonates took 24–30 more hours to mature in the presence of copepods than in the absence of copepods. Enlarged body and spine lengths of progeny coincided with reduced progeny number during the first six reproductive instars. The disadvantage of the reduced offspring number produced per female was balanced by enhanced survivorship of progeny subjected to copepod predation.     

Observations on the effect of photoperiod on the control of polymorphism in Psylla pyricola

ABSTRACT. Psylla pyricola Forster (Homoptera) occurs as two adult morphs, a summer form (f. typica) and a winter form (f. simulans). The latter is larger, darker and overwinters in ovarian diapause. Eggs laid by either morph under LD 12:12 (short days) or under LD 18:6 or 16:8 (long days) gave f. typica when reared under long days. When reared under short days, almost all were f. simulans. No interforms were recorded. When the photoperiod was switched during the nymphal stage (whether from long to short days, diapause induction; or from short to long days, diapause termination), nymphs were progressively less sensitive with increasing age, with no morph changes when the switch occurred in the fifth instar. Nymphs derived from f. simulans reared under long days tended to be more sensitive to a change to short days in the early instars than when the nymphs were first reared under short days and then switched to long days. Interforms were frequent when the switch occurred during the nymphal stage, particularly in the third instar. Wing lengths varied within morphs depending on the timing of the switch. The largest f. simulans were derived from f. typica when maintained under short days, and the smallest f. typica from either morph when maintained under long days, as would be expected under field conditions. No f. simulans were found under field conditions in the first f. typica generation, but a small percentage were present in the second generation. It is concluded that, even within a given generation, not all P. pyricola are equally sensitive to photoperiod, but that photoperiod is one of the major factors controlling the change to f. simulans in the autumn. Whilst under laboratory conditions photoperiod can also affect the switch to f. typica in the spring, in the field it may be less important than low temperatures.     

Seasonal polyphenism of spotted‐wing Drosophila is affected by variation in local abiotic conditions within its invaded range,likely influencing survival and regional population dynamics

1. Overwintering Drosophila often display adaptive phenotypic differences beneficial for survival at low temperatures. However, it is unclear which morphological traits are the best estimators of abiotic conditions, how those traits are correlated with functional outcomes in cold tolerance, and whether there are regional differences in trait expression.
2. We used a combination of controlled laboratory assays, and collaborative field collections of invasive Drosophila suzukii in different areas of the United States, to study the factors affecting phenotype variability of this temperate fruit pest now found globally.
3. Laboratory studies demonstrated that winter morph (WM) trait expression is continuous within the developmental temperature niche of this species (10–25°C) and that wing length and abdominal melanization are the best predictors of the larval abiotic environment.
4. However, the duration and timing of cold exposure also produced significant variation in development time, morphology, and survival at cold temperatures. During a stress test assay conducted at ?5°C, although cold tolerance was greater among WM flies, long‐term exposure to cold temperatures as adults significantly improved summer morph (SM) survival, indicating that these traits are not controlled by a single mechanism.
5. Among wild D. suzukii populations, we found that regional variation in abiotic conditions differentially affects the expression of morphological traits, although further research is needed to determine whether these differences are genetic or environmental in origin and whether thermal susceptibility thresholds differ among populations within its invaded range.

     

Parasitoids induce production of the dispersal morph of the pea aphid, Acyrthosiphon pisum

In animals, inducible morphological defences against natural enemies mostly involve structures that are protective or make the individual invulnerable to future attack. In the majority of such examples, predators are the selecting agent while examples involving parasites are much less common. Aphids produce a winged dispersal morph under adverse conditions, such as crowding or poor plant quality. It has recently been demonstrated that pea aphids, Acyrthosiphon pisum, also produce winged offspring when exposed to predatory ladybirds, the first example of an enemy‐induced morphological change facilitating dispersal. We examined the response of A. pisum to another important natural enemy, the parasitoid Aphidius ervi, in two sets of experiments. In the first set of experiments, two aphid clones both produced the highest proportion of winged offspring when exposed as colonies on plants to parasitoid females. In all cases, aphids exposed to male parasitoids produced a higher mean proportion of winged offspring than controls, but not significantly so. Aphid disturbance by parasitoids was greatest in female treatments, much less in male treatments and least in controls, tending to match the pattern of winged offspring production. In a second set of experiments, directly parasitised aphids produced no greater proportion of winged offspring than unparasitised controls, thus being parasitised itself is not used by aphids for induction of the winged morph. The induction of wing development by parasitoids shows that host defences against parasites may also include an increased rate of dispersal away from infected habitats. While previous work has shown that parasitism suppresses wing development in parasitised individuals, our experiments are the first to demonstrate a more indirect influence of parasites on insect polyphenism. Because predators and parasites differ fundamentally in a variety of attributes, our finding suggests that the wing production in response to natural enemies is of general occurrence in A. pisum and, perhaps, in other aphids.     

Seasonal and reciprocal succession and cyclomorphosis of two Bosmina species (Cladocera, Crustacea) co-existing in a lake: their relationship with invertebrate predators

The seasonal succession and cyclomorphosis of Bosmina longirostrisand Bosmina fatalis (also known as Sinobosmina fatalis) wereinvestigated in the shallow eutrophic Lake Suwa from 1996 to1999. Two different invertebrate predators, cyclopoid copepodsand Leptodora kindtii, co-existed with Bosmina in the lake.A pattern of seasonal succession of the two species was apparent,with B. fatalis dominant in summer and B. longirostris in springand fall. The two species also showed different seasonal changesin morphology. During summer, when the density of the invertebratepredator Leptodora was high, B. fatalis changed its morphologyand maintained long features, with elongation of the antennulesand mucrones. This species also maintained the protruding shapeof the antennules by increasing the angles between the antennulesand the body, and between two antennules, during the same period.The seasonal differences and magnitude of changes in morphologywere more marked for larger individuals. However, such markeddevelopment of protruding structures was not observed in B.longirostris, which showed only slight changes in the antennuleand mucrone lengths related to the density of copepods. Theresults suggest that the morphological traits of B. fatalisare a morphological defence against Leptodora, while those ofB. longirostris are a response to copepods. In Lake Suwa, thepredators show different patterns of seasonal changes in density.Different morphological traits and the resulting interactionswith invertebrate predators seem to be important in determiningthe reciprocal succession of the two Bosmina species.     

Thermal biology of the common garter snake Thamnophis sirtalis (L.)

Summary The thermoregulatory significance of a striped-melanic colour polymorphism in the common garter snake, Thamnophis sirtalis, was assessed through a combination of labortory experimentation and field study. In experiments with living snakes the melanic morph maintained a higher body temperature than the striped morph, when exposed to natural insolation. Experiments with excised skin showed that this thermal advantage is attributable to some integumental difference between the two morphs. Body temperatures of snakes in the field revealed that, during the colder part of the active season, melanics were able to stay warmer than striped snakes by an amount (1.24 C°) approximating the difference observed in the laboratory. Some evidence and argument is presented to suggest that melanism also may confer protection against overheating in warm periods.     

Expression of the heat-shock protein HSP70 in Drosophila buzzatii lines selected for thermal resistance

The level of HSP70 expression induced by a non-lethal high temperature was examined in lines selected for increased thermal resistance and in corresponding control lines of Drosophila buzzatii, in order to test if selection for high temperature resistance leads to an increased level of HSP70 expression. The lines used were selected for up to 64 generations either as adults or through all larval stages. In adult selection lines, hard selection was implemented every second generation after mild heat hardening. In larval selection lines, larvae were exposed each generation to laboratory "natural" selection. Generally lines selected as adults showed a higher HSP70 expression than did controls, both in third instar larvae and in adults. A strong negative response to selection of HSP70 expression was found in all lines that were selected at cycling temperatures during larval development. The results suggests that a trade off between heat resistance in form of HSP70 expression and fecundity/fertility are responsible for the level of HSP70 expression. The effect of the different methods of selection on HSP70 expression suggests that heat resistance constitutes more than one trait.     

Morphological variation in Bosmina longirostris (O.F. Müller, 1785) (Crustacea: Cladocera): consequence of cyclomorphosis or indication of cryptic species?

Samples of the polymorphic cladoceran Bosmina longirostris were collected in several lakes and ponds in Western Germany between 1993 and 1999. The shape of the first antennule was used as a qualitative criterion to classify 1241 individuals to three morphs (`longirostris', `cornuta', `pellucida') according to Lieder (Crustacea: Cladocera: Bosminidae. Stuttgart, Germany: Gustav. Fischer Verlag, 1996). Eight morphometric variables were subsequently measured in each individual to quantify size‐ and shape‐related features. Principal component and discriminant analyses were used to describe the morphological variation within and among the three morphs. Seasonal environmental changes modified several traits and thus, cyclomorphosis accounted for a high proportion of within‐morph variation. However, there was no gradual change from one morph to another. As well among‐morph variation was considerably greater than within‐morph variation. Consequently, the disagreement between the qualitative pre‐classification and the assignment basing on morphometric discriminant functions was low (4.6–12.0%). Considering that each morph is morphologically well‐defined, and that the different morphs coexisted over several generations in the same lake we conclude that they represent different species rather than variants of one polymorphic species. Therefore, we recognize beside the nominal taxon Bosmina longirostris ( 26 ) two more species: Bosmina cornuta ( 10 ) and Bosmina pellucida 30 .     

An interaction‐driven cannibalistic reaction norm

Cannibalism is induced in larval‐stage populations of the Hokkaido salamander, Hynobius retardatus, under the control of a cannibalism reaction norm. Here, I examined phenotypic expression under the cannibalism reaction norm, and how the induction of a cannibalistic morph under the norm leads to populational morphological diversification. I conducted a set of experiments in which density was manipulated to be either low or high. In the high‐density treatment, the populations become dimorphic with some individuals developing into the cannibal morph type. I performed an exploratory analysis based on geometric morphometrics and showed that shape characteristics differed between not only cannibal and noncannibal morph types in the high‐density treatment but also between those morph types and the solitary morph type in the low‐density treatment. Size and shape of cannibal and noncannibal individuals were found to be located at either end of a continuum of expression following a unique size–shape integration rule that was different from the rule governing the size and shape variations of the solitary morph type. This result implies that the high‐density‐driven inducible morphology of an individual is governed by a common integration rule during the development of dimorphism under the control of the cannibalism reaction norm. Phenotypic expression under the cannibalism reaction norm is driven not only by population density but also by social interactions among the members of a population: variation in the populational expression of dimorphism is associated with contingent social interaction events among population members. The induced cannibalistic morph thus reflects not only by contest‐type exploitative competition but also interference competition.     

Mechanical vibrations from tadpoles' flapping tails transform salamander's carnivorous morphology

Some prey or predator organisms exhibit striking rapid morphological plastic changes with distinct morphology under the condition of predator or prey presence. Remote chemicals propagating from the inducing agents are the prevalent induction cues for most examples of induction of distinct morphs. Sonic and visual cues, as well as chemical cues, are known as triggers for induction of behavioural plasticity. Here we show that hydraulic vibration originating from flapping tails of anuran tadpoles is a key cue in relation to induction of a distinct carnivorous morphology, a broad-headed morph, in larval salamander Hynobius retardatus, which is able to efficiently capture and handle prey. This result was further supported by the fact that simple mechanical vibrations of tail-like vinyl fins were able to induce the morph without any biological cues. Induction of the morph triggered by hydraulic vibration provides a novel concept for understanding the proximate mechanisms of induction of morphological changes.     

Alternative life histories in the water strider Gerris lacustris: time constraint on wing morph and voltinism

The wing dimorphic water strider Gerris lacustris L. (Heteroptera: Gerridae) switches to a bivoltine life cycle under favorable climatic conditions. The switch in voltinism is accompanied by a reduction of wing development in the directly reproducing midsummer generation, while the diapausing generation has a high fraction of long‐winged individuals. We investigated whether the thermal energy (degree‐days) available in natural habitats constrains the combination of developmental pathway and wing morph. Offspring of G. lacustris were reared under quasi‐natural conditions at two temperature regimes to determine the thermal constant k required to complete adult development in either wing morph. The thermal constant for egg‐to‐adult development of the short‐winged morph was about 20% lower than of the long‐winged morph. Based on the results from the outdoor laboratory experiment, we calculated the total degree‐days necessary to complete the possible combinations of wing morph pattern and voltinism. Comparison of these estimates with the thermal energy actually available during the reproductive season of 2004 for various natural habitats (sun‐exposed field ponds and shaded forest ponds) suggests that voltinism as well as wing morph pattern is strongly limited by the number of degree‐days available in these habitats. On forest ponds, only univoltine life cycles were possible, whereas on field ponds temperature allowed bivoltine life cycles. However, only the eggs laid at the very beginning of the season had the potential to accumulate enough degree‐days to complete a bivoltine life cycle with both generations long‐winged. We conclude that thermal energy is the main environmental constraint limiting voltinism of populations in the two habitat types. Furthermore, the available thermal energy also seems to influence the determination of the seasonal wing pattern in G. lacustris.     

Role of photoperiod and temperature in seasonal morph determination of the butterfly Eurema hecabe

ABSTRACT. Environmental cues affecting seasonal morph determination of the pierid butterfly Eurema hecabe L. were examined in laboratory experiments and in the field. A sexual difference exists in the photoperiodic response for seasonal morph determination: in short days at 25C the proportion of the autumn morph is higher in females than in males, and as the rearing temperature is decreased the proportion of autumn morphs increases in both sexes. A 5C drop in temperature during the final larval stadium induces 100% autumn morph production in short-day animals. This increase in autumn morph production induced by temperature decrease explains the seasonal morph change of this butterfly occurring in early autumn in the field. The role of this sex-linked photoperiodic response is discussed as a seasonal strategy of reproduction.     

异质生境对二型花柱植物雪地报春花性状分化的影响

该研究以生长在新疆北部的二型花柱植物雪地报春(Primula nivalis)为研究材料,对7个低海拔(1 657~2 013m)森林居群和7个高海拔(2 400~2 700m)草原居群的花型比例进行调查,对花部特征进行检测,并对长、短花柱植株在不同海拔居群的花性状分化进行了比较研究,以揭示该类群植物的演化与环境的关系。结果表明:(1)在高海拔草原居群中长花柱植株比例高于短花柱植株。(2)比较草原和森林生态系统中的雪地报春居群,长花柱花形态特征间存在显著差异,但短花柱花形态特征间的差异不显著。(3)高海拔草原居群中长花柱花的花冠筒长度、雄蕊长度、雌雄蕊距离、雌雄异位程度以及雌雄蕊空间位置是导致花性状分化的主要因素。(4)在草原居群中,长花柱花的雄蕊、雌雄异位程度及花冠口至雄蕊距离的变化减少了长短花柱花间的互补式对应性。研究认为,在异质生境中长花柱花性状的变化是导致该物种长短花柱花性状分化的主要因素。     

Aphid Wing Induction and Ecological Costs of Alarm Pheromone Emission under Field Conditions

The pea aphid, Acyrthosiphon pisum Harris, (Homoptera: Aphididae) releases the volatile sesquiterpene (E)-β-farnesene (EBF) when attacked by a predator, triggering escape responses in the aphid colony. Recently, it was shown that this alarm pheromone also mediates the production of the winged dispersal morph under laboratory conditions. The present work tested the wing-inducing effect of EBF under field conditions. Aphid colonies were exposed to two treatments (control and EBF) and tested in two different environmental conditions (field and laboratory). As in previous experiments aphids produced higher proportion of winged morphs among their offspring when exposed to EBF in the laboratory but even under field conditions the proportion of winged offspring was higher after EBF application (6.84±0.98%) compared to the hexane control (1.54±0.25%). In the field, the proportion of adult aphids found on the plant at the end of the experiment was lower in the EBF treatment (58.1±5.5%) than in the control (66.9±4.6%), in contrast to the climate chamber test where the numbers of adult aphids found on the plant at the end of the experiment were, in both treatments, similar to the numbers put on the plant initially. Our results show that the role of EBF in aphid wing induction is also apparent under field conditions and they may indicate a potential cost of EBF emission. They also emphasize the importance of investigating the ecological role of induced defences under field conditions.