Life history plasticity of Nemoura trispinosa (Plecoptera: Nemouridae) along a permanent-temporary water habitat gradient

1. The life history of the small herbivorous stonefly Nemoura trispinosa Claassen was studied in a variety of small springs in southern Ontario, Canada. Nymphs generally were able to tolerate a wide range of environmental conditions and were found in 78% of habitats sampled, although population densities differed markedly. 2. Life-cycle patterns varied from a univoltine, slow seasonal type to a univoltine, fast seasonal type with extended egg development. In one, highly stable, spring the life cycle was semivoltine. Inter-year variation was studied for 5 years in one spring and was found to be low relative to among-spring variation. 3. Differences in the life history traits of N. trispinosa populations from our spring series were most probably an expression of phenotypic plasticity rather than of genetic differentiation. 4. Maximum annual water temperature was the factor most influential on nymphal growth rate (non-linear relationship), whereas range in generation time was related to the degree of habitat permanence.     

Biology and damage traits of emerald ash borer planipennis Fairmaire）in China

Emerald ash borer （Agrilus planipennis Fairmaire） （Coleoptera： Buprestidae） is a major stem borer of ash （Fraxinus spp.）. It is univoltine in Tianjin, while it is semivoltine in Heilongjiang Province, and both univoltine and semivoltine in Changchun, Jilin Province, where the majority is univoltine. The longevity of emerald ash borer adults is 17.2 ± 4.6 days （n = 45）, eggs 9.0 5：1.1 days （n = 103）, univoltine larvae 308 days, semivoltine larvae 673 days, and pupae 61.2 ± 1.6 days （n = 45）. It takes about 100 days from the time larvae bore into the phloem to when they complete the pupal cell. In a 10-year-old velvet ash （Fraxinus velutina Tort.） plantation in Tianjin, emerald ash borer preferred to oviposit on the regions of boles from 50-150 cm above ground, accounting for 76.7% of the total girdling. Girdling on the south side of the tree boles accounted for 43.40% of the total girdling. The emerald ash borer population density is higher at the edge of the plantation compared with the center.     

Live cycles,drift, and standing stocks of some stoneflies (Insecta: Plecoptera) from streams in Minnesota,U.S.A.

From analyses of monthly length frequency distributions of stonefly (Plecoptera) nymphs from stream riffles, two types of univoltine life cycles (Hynes 1961, classification) were observed: slow seasonal forIsoperla transmarina, Isoperla signata, Isogenoides olivaceus, andAllocapnia rickerii, and fast seasonal forProstoia similis. The Perlodidae stoneflies (Isoperla andIsogenoides) showed greatest propensities to drift near the end of their life cycles. Drift rates of the other two species of stoneflies were correlated to standing stock estimates.     

Resource dynamics and detritivore production in an acid stream

1. Life history patterns and production of eight shredder‐detritivore species were studied in relation to the detritus dynamics of a small acidic stream in England. Mean annual detritus inputs (direct and lateral sources combined) were approximately 400 g m^?2 year^?1 and showed significant seasonal and annual variation. 2. Detritus standing stock did not increase significantly during times of high input, reflecting low retention efficiency. However, the mean detritus standing stock was relatively large (108 g m^?2) reflecting a slow decomposition rate typical of acid streams. 3. Four species were univoltine with highly synchronous patterns of emergence and recruitment (Leuctra inermis, Leuctra hippopus, Capnia vidua and Amphinemura sulcicollis). Two species were univoltine with extended patterns of emergence and recruitment (Nemoura cinerea, Potamophylax cingulatus). Leuctra nigra was apparently semivoltine, while Protonemura meyeri showed two successive cohorts in the second year of the study, suggesting either bivoltinism or cohort splitting. 3. Secondary production of the dominant shredders was 1.67 g m^?2 year^?1 in 1997 and 1.99 g m^?2 year^?1 in 1998, which is low compared with other small European streams. This was probably because of an impoverished invertebrate community and poor food quality associated with acid conditions. Food availability probably did not account for the low production as the detritus standing stock far exceeded the estimated shredder ingestion of 42–50 g m^?2 year^?1. 4. Despite low overall shredder production, species‐specific production was high, possibly because of competitive release in this species‐poor acid stream. Periods of high production and growth showed no relationship with detritus availability but were closely related to life history.     

Life History of a Population of Protonemura intricata (Ris, 1902) (Insecta,Plecoptera) in a Constant Temperature Stream in Central Europe

The nymphal biology (life cycle, secondary production and feeding) of a population of Protonemura intricata was studied in a calcareous stream with almost constant temperature (7–8 °C) through the year at Prosiek valley (Chočské Vrchy Mts., West Carpathians, Slovakia). The results were compared with those previously known for this species in other areas. According to our data, the species has a complicated life cycle consisting of two cohorts with different duration (probably a cohort splitting). This life cycle seems to be uncommon among the Plecoptera, which usually has simple univoltine or semivoltine life cycles, and could be probably attributed to the constant temperature regime similar to that of tropical environments. One factor, the photoperiod length, was positively correlated with monthly growth rates. The nymphal growth varied throughout the year, but the highest growth was in late spring and summer, and the lowest was in winter. The annual secondary production was very high (909.4 mg · m^–2) despite not being the most frequent taxon in the stream. The highest monthly production values were observed in the autumnal‐winter period (November–December) due to massive hatching of nymphs. The lowest production values were observed in late summer and early autumn. Analyses of gut contents showed that this species behaves mainly as gatherer‐collector, but also as shredder and, occasionally, as scraper. Changes in the gut content composition were detected in relation to the size. Comparing our results with those of the literature, it can be inferred that different populations of P. intricata can show notable differences in their life history when inhabiting places with different thermal conditions. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Long‐term monitoring of the life history of a riverine mayfly Ephemera orientalis (Ephemeroptera: Ephemeridae) in the Han River in Seoul,Korea

Climatic change often affects life history aspects of aquatic insects. Long‐term monitoring was conducted to understand the life history variability of the riverine mayfly Ephemera orientalis in the Han river, where habitat is largely disturbed by summer floods. Water level of the study site, Godeok‐dong area in Seoul, is regulated by the Paldang Dam located approximately 14 km above the study site. E. orientalis often emerged in large numbers around the study area. Larvae were sampled monthly from April 2006 to July 2010 using a Surber sampler (50 × 50 cm, mesh 0.25 mm, two replicates). Additional qualitative sampling was conducted using a hand net. As a result, based on the larval body size distribution, E. orientalis had a univoltine life cycle with two distinct slow growing cohort groups: the S1group emerged during May–June and the S2 group emerged during August–September. Our previous study conducted in Gapyeong stream in Gyeonggi‐do showed that E. orientalis normally had three cohort groups, S‐1 and S‐2 groups and the F‐group (a fast growing cohort developed during May–August). However, the Han River population of E. orientalis lacked such a fast growing cohort. It is most probable that habitat disturbances caused by high water level during the rainy season negatively affected the development of the fast growing cohort (F‐group) in the Han River area. The relationships between monthly E. orientalis population data and water level in the Han River are presented.     

Life histories,abundance and distribution of some macroinvertebrates from a South Carolina,USA coastal plain stream

The life histories, densities and distribution patterns of the most abundant macroinvertebrates, exclusive of chironomids and oligochaetes, are reported for a low-gradient, second-order, blackwater stream on the Coastal Plain of South Carolina, USA. Univoltine life histories were found for all species of Coleoptera (Ancyronyx variegata, Macronychus glabratus, Dubiraphia quadrinotata and Hydroporus clypealis) and most species of Ephemeroptera (Eurylophella temporalis, Paraleptophlebia volitans and Hexagenia munda), although the mayfly Stenonema modestum was at least bivoltine. Both univoltinism (Macrostemum carolina and Pycnopsyche luculenta), partial bivoltinism (Nyctiophylax affinis) and complete bivoltinism (Hydropsyche decalda, Cheumatopsyche sp., Phylocentropus placidus and probably Chimarra florida) were found among the Trichoptera. Odonate species were both univoltine (Calopteryx dimidiata, Enallagma divagans and Epitheca cynosura) and semivoltine (Boyeria vinosa, Gomphus lividus and Macromia georgina). The alderfly Sialis aequalis and the isopod Asellus laticaudatus were univoltine, while the blackfly Simulium taxodium produced at least 6 generations per year. Groups of functionally-related species within the orders Odonata, Trichoptera and Coleoptera each exhibited possible temporal and/or spatial segregation.     

Diapause and overwintering of two spruce bark beetle species

Diapause, a strategy to endure unfavourable conditions (e.g. cold winters) is commonly found in ectothermic organisms and is characterized by an arrest of development and reproduction, a reduction of metabolic rate, and an increased resistance to adversity. Diapause, in addition to adaptations for surviving low winter temperatures, significantly influences phenology, voltinism and ultimately population growth. We review the literature on diapause and overwintering behaviour of two bark beetle species that affect spruce‐dominated forests in the northern hemisphere, and describe and compare how these strategies can influence population dynamics. The European spruce bark beetle Ips typographus (L.) (Coleoptera, Curculionidae) is the most important forest pest of Norway spruce in Europe. It enters an adult reproductive diapause that might be either facultative or obligate. Obligate diapausing beetles are considered strictly univoltine, entering this dormancy type regardless of environmental cues. Facultative diapausing individuals enter diapause induced by photoperiod, modified by temperature, thus being potentially multivoltine. The spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) infests all spruce species in its natural range in North America. A facultative prepupal diapause is averted by relatively warm temperatures, resulting in a univoltine life cycle, whereas cool temperatures induce prepupal diapause leading to a semivoltine cycle. An adult obligate diapause in D. rufipennis could limit bi‐ or multivoltinism. We discuss and compare the influence of diapause and overwinter survival on voltinism and population dynamics of these two species in a changing climate and provide an outlook on future research.     

The effect of temperature on the egg incubation period and nymphal growth of two Nemoura species (Plecoptera) from subarctic fennoscandia

Field studies of Nemoura arctica and N. viki showed that the two species preferred different biotopes and indicated differences in their life‐cycles. N. arctica seemed to have a semivoltine life‐cycle, at least in some years, this was not the case for N. viki.

Laboratory studies showed that the temperature tolerance of the eggs of N. arctica was wider than for those of N. viki. The length of the egg incubation period of both species was influenced by the ambient temperature, but no significant interspecific difference existed between the regression lines of the relationship between the temperature (T°C) and egg incubation period (Y days), as given by the regression equation Y = aT^?b for the log values. Reared at constant food supply, nymphal growth occurred in two periods. Firstly a rapid growth to about 4 mm, followed by a period of slow growth until emergence. During the first period growth (G)at (D) days was linear, according to the simple linear regression equation G = a+bD . The temperature tolerance of the nymphs of N. arctica was wider than for those of N. viki and significant interspecific differences between the species in growth were recorded at relatively high rearing temperatures, such as 12° and 16°C, but not at low temperatures. At constant food supply, nymphal growth was greatly influenced by the rearing temperature. At a mean temperature of 16°C N. arctica nymphs grew rapidly and emerged after 120 days; whereas at a mean temperature of 4°C growt h was slow and the nymphs did not even manage to reach the emergence stage after 700 days.     

Life cycle and nymphal feeding of Rhabdiopteryx christinae Theischinger 1975 (Plecoptera: Taeniopterygidae)

In the present paper some aspects of the nymphal biology of Rhabdiopteryx christinae, an endemic species from the Iberian Peninsula, are studied in a temporary stream from Southern Spain. R. christinae has an univoltine life cycle, with very short, fast and constant nymphal development, only having four months (December-March), related with the seasonality of the stream. In order to resist the period in which the stream has no water, an egg diapause probably occurs. The study of nymphal gut contents showed that R. christinae acts mainly as collector (detritus) and scraper (epiphytic and epilitic diatoms), as it is generally accepted for the family Taeniopterygidae.     

The management of metabolic energy storage during the life cycle of mayflies: a comparative field investigation of the collector-gatherer <Emphasis Type="Italic">Ephemera danica</Emphasis> and the scraper <Emphasis Type="Italic">Rhithrogena semicolorata</Emphasis>

The concentration and seasonal dynamics of the major energy storage components, triglycerides and glycogen, were measured in two species of mayfly (Rhithrogena semicolorata and Ephemera danica) with contrasting life cycle strategies living in a small mountain stream. E. danica is a burrowing, semivoltine collector-gatherer; R. semicolorata is univoltine and scrapes periphyton from stones. This is the first publication which focuses on the role of metabolic energy sources during the larval life span of two mayfly species until the larvae emerge. Although triglycerides are the major energy reserve in both species (>84% of total energy storage) throughout the whole larval development their seasonal dynamic differed considerably. In R. semicolorata the triglyceride concentration declined during the last weeks prior to emergence in both sexes. The same pattern was found in female larvae of E. danica, but not in male E. danica. It is suggested that females use triglycerides in the last larval stages for egg maturation, which is completed in the last larval instar. In male E. danica the triglyceride concentrations remained high until emergence, presumably due to their high energy demands as adults for their swarming flights. Glycogen concentrations did not show such a difference between species and sexes. Its significance as a storage substrate for energy is rather low; however, concentrations decreased in both species and sexes prior to emergence.     

Temperature-induced changes in the life cycle of Leuctra nigra(Plecoptera: Leuctridae) from a Lake District stream

1. The life cycle of Leuctra nigra (Olivier) took 2 years in a small stream in the English Lake District and the exponential growth of the larvae was scarcely affected by variations in water temperature (range 4.2-14.0°C). Mean growth rates for three year-classes were 0.43±0.01, 0.42±0.01, 0.39±0.05% body length day^?1. There were thirteen or fourteen larval instars for males and fourteen or fifteen for females. The ratio between successive instars was a constant 1.20 (conformed with Dyar's rule). 2. Larval growth and mortality were exponential at six constant temperatures (5.9, 8.2, 12.1, 15.8, 18.2, 19.8°C) in the laboratory. Mean growth rates (% body length day^?1) increased directly with temperature from 0.37 (5.9°C) to 0.55 (19.8°C). Mean mortality rates (% day^?1) increased directly with temperature from 0.20 (5.9°C) to 0.26 (12.1°C) and then markedly increased to 0.54-0.58 at the three higher temperatures. Only 7-10% of animals completed their life cycle at the three higher temperatures compared with 23–27% at the three lower temperatures. Egg production also decreased considerably at the higher temperatures. 3. As growth rates in the stream and laboratory were similar at similar temperatures (<14°C), the optimum conditions for growth in the laboratory were probably similar to those in the stream; therefore resources such as food and space were not restricting growth in the stream. 4. The implications of the temperature-induced changes in growth and mortality are discussed and it is concluded that although the life cycle can change from semivoltine to univoltine with increasing temperature, the costs of a univoltine life cycle are high in terms of survival and egg production, both of which decreased markedly between 12.1 and 15.8°C. Therefore the optimum habitat for this species appears to be a summer cool stream (maximum temperature <14°C) and the optimal life cycle appears to be about 2 years from egg to adult.     

Mayfly production in a Colorado mountain stream: an assessment of methods for synchronous and non-synchronous species

Year-round collections of mayflies (Ephemeroptera) from a Colorado mountain stream allowed critical examination of several methods of calculating production for species with different life cycles. Five of the six numerically dominant species exhibited slow seasonal, univoltine life cycles. Baetis tricaudatus was bivoltine. Two species demonstrated well synchronized development, three species were poorly synchronized and a sixth was intermediate. Mean density and biomass data from each sampling date were used to ascertain the goodness-of-fit of each species to the Allen curve. It is proposed that such information can provide quantitative criteria for identifying species with well synchronized development and thereby determine when it is appropriate to directly apply cohort methods while avoiding time intensive body size (e.g. head width) measurements necessary for size-frequency analyses. In addition, these data demonstrate that species specific production varies with gross changes in elevation.     

Integrating life history and physiology to understand latitudinal size variation in a damselfly

Our understanding of latitudinal life history patterns may benefit by jointly considering age and mass at maturity and growth rate. Additional insight may be gained by exploring potential constraints through pushing growth rates to their maximum and scoring physiological cost‐related variables. Therefore, we reared animals of a univoltine Spanish and Belgian population and of a semivoltine Swedish population of the damselfly Enallagma cyathigerum (spanning a latitude gradient of ca 2350 km) in a common environment from the eggs until adult emergence and exposed them to a transient starvation period to induce compensatory growth. Besides age and mass at maturity and growth rate we also scored investment in energy storage (i.e. triglycerides) and immune function (i.e. total activity of phenoloxidase). At emergence, body mass was greater in Spain and Sweden and lower in Belgium, suggesting a genetic component for the U‐shaped latitudinal pattern that was found also in a previous study based on field‐collected adults. The mass difference between univoltine populations can be explained by the shorter development time in the Belgian population, and this despite a higher growth rate, a pattern consistent with undercompensating countergradient variation. In line with the assumed shorter growth seasons, Belgian and Swedish animals showed higher routine growth rates and compensatory growth after transient starvation. Despite a strong link with metabolic rates (as measured by oxygen consumption) populations with higher routine growth rates had no lower fat content and had higher immune function (i.e. immune function decreased from Sweden to Spain), which was unexpected. Rapid compensatory growth did, however, result in a lowered immune function. This may contribute to the absence of perfect compensating countergradient variation in the Belgian population and the lowest routine growth rates in the Spanish population. Our results underscore the importance of integrating key life historical with physiological traits for understanding latitudinal population differentiation.     

Termination of pupal diapause in the pine processionary moth Thaumetopoea pityocampa

Diapause development is a complex process involving several eco‐physiological phases. Understanding these phases, especially diapause termination, is vital for interpreting the life history of many insect species and for developing suitable predictive models of population dynamics. The pine processionary moth is a major defoliator of pine and a vertebrate health hazard in the Mediterranean region. This species can display either univoltine or semivoltine development, with a pupal diapause extending from a few months to several years, respectively. Although the ecological and applied importance of diapause is acknowledged, its physiological regulation in either case remains obscure. In the present study, we characterize pre‐termination, termination and post‐termination phases of pupae developing as univoltine or remaining in prolonged diapause. Changes in metabolic activity are monitored continuously using thermocouples, comprising a novel method based on direct calorimetry, and periodically by use of O₂ respirometry. The two methods clearly detect diapause termination in both types of pupae before any visible morphological or behavioural changes can be observed. Univoltine individuals are characterized by an increase in metabolic activity from pre‐termination through to termination and post‐termination, ultimately resulting in emergence. Remarkably, a synchronous termination is observed in individuals that enter prolonged diapause instead of emerging; however, in these pupae, the increased metabolic activity is only transient. The present study represents a starting point toward understanding the eco‐physiology of diapause development processes in the pupae of the pine processionary moth.     

Instar sizes,life cycles,and food habits of five Rhyacophila (Trichoptera: Rhyacophilidae) species from the Appalachian Mountains of South Carolina,U.S.A.

The larval head widths at each instar, life cycles, and food habits of late instars were determined for five species of Rhyacophila from two Appalachian mountain streams in South Carolina, U.S.A. Rhyacophila acutiloba Morse & Ross was univoltine with two cohorts, one emerging in the spring and another presumably emerging in early autumn. Rhyacophila fuscula (Walker), R. nigrita Banks, and R. carolina Banks were apparently multicohort, univoltine species with extended flight periods. Rhyacophila minor Banks was univoltine with a spring emergence. All species were predaceous and consumed mainly Plecoptera nymphs and Trichoptera larvae.     

Mating disruption of pea moth (Cydia nigricana) in organic peas (Pisum sativum)

We appraised mating disruption (MD) to control pea moth, Cydia nigricana (Fabricius) (Lepidoptera: Tortricidae), by assessing male attraction to monitor traps, larval pod infestation, and larval age structure in pheromone‐treated and untreated grain pea fields [Pisum sativum L. (Fabaceae)], over a 5‐year period. Cellulose pheromone dispensers were manually attached to the top shoots of pea plants and released 540 mg ha^?1 day^?1 synthetic pheromone E8,E10‐dodecadien‐1‐yl acetate in a first test series (2000–2001) and ca. 4 200 mg pheromone ha^?1 day^?1 in a second series (2004–2006). The dispensers had a half‐life of about 30 days. Although male attraction to pheromone monitoring traps was largely suppressed at the edges and within MD fields in both test series, MD treatments did not reduce pod infestation in the open field in 2000 and 2001. In the 2004–2006 series, larval damage reduction was achieved in the majority of the trials but overall MD efficacy in the open field was only 61% and not significant. In contrast, in field cages placed within the experimental sites and supplied with unmated pea moths, MD control was consistently high and significant. There were no obvious differences in the larval age distribution in all MD and control treatments, suggesting that infestations started and developed further similarly. As a univoltine species, C. nigricana larvae stay in the soil of pea fields for hibernation and pupate. The following year, emerging adults disperse and fly to the closest pea crop. Combined emergence site and pea crop treatments were conducted over 2 years to include this early migration phase of C. nigricana adults. However, the emergence site treatments did not enhance MD‐control efficacy. We conclude that mating activity was only prevented in cage tests, whereas substantial mating occurred during the transit phase outside the pheromone‐treated fields either within non‐crop vegetation and/or at the edges of pheromone‐treated pea fields orientated upwind. Thus, resulting gravid female entry can be regarded as the major constraint to reliable MD control.     

Temperature‐Dependent Growth and Life Cycle of Nemoura sichuanensis (Plecoptera: Nemouridae) in a Chinese Mountain Stream

Plecoptera constitute a numerically and ecologically significant component in mountain streams all over the world, but little is known of their life cycles in Asia. The life cycle of Nemoura sichuanensis and its relationship to water temperature was investigated during a 4‐year study in a headwater stream (known as the Jiuchong torrent) of the Xiangxi River in Central China. Size structure histograms suggest that the life cycle was univoltine, and the relationships between the growth of Nemoura sichuanensis, physiological time, and effective accumulated water temperature were described using logistic regressions. The growth pattern was generally similar within year classes but growth rates did vary between year‐classes. Our field data suggest a critical thermal threshold for emergence in Nemoura sichuanensis, that was close to 9 °C. The total number of physiological days required for completing larval development was 250 days. The effective accumulated water temperature was 2500 degree‐days in the field. Development during the life cycle increased somewhat linearly with the physiological time and the effective accumulated water temperature, but some non‐linear relationships were best developed by logistic equations. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Nymphal biology of Ephoron virgo (Olivier, 1791) (Ephemeroptera,Polymitarcyidae) in an Apenninic river (NW Italy)

Life cycle, feeding habits and nymphal density of a population of Ephoron virgo from the Bormida river (Northwestern Italy) were studied. Nymphs were present in the river from the beginning of May until August, with the highest density of nymphs recorded at the end of May–beginning of June. The life cycle was univoltine and the nymphal growth was fast. During the life cycle, an obligatory egg diapause occurs and individuals remain in this stage for up to nine months. Nymphs fed mainly on detritus, although mineral matter constituted a considerable percentage of the gut contents, predominantly in smaller nymphs.     

Life history variability of a grazing stream insect (Liponeura cinerascens minor; Diptera: Blephariceridae)

SUMMARY 1. Fourteen populations of Liponeura cinerascens minor from different sites in the Swiss Alps were investigated. Our goals were to describe the life history of this blepharicerid species and to analyse how it was influenced by temperature and food availability. 2. Temperature regimes and periphyton density at the sampling sites varied considerably. Mean annual temperature ranged from 3.8 to 6.5 °C, with annual amplitudes (i.e. the difference between the mean of the coldest and the warmest month) between 3.7 and 12.0 °C. Averaged periphyton density, which was assessed semiquantitatively on a scale from 0 to 5, ranged from 1.1 to 3.3. 3. Larvae or pupae of L. c. minor were found between January and October. Presence of larvae or pupae at individual sites ranged from 100 to 224 days and appeared to be mainly a result of hatching patterns. Individual larval development required between 7 and 20 weeks. Our findings suggest that L. c. minor is univoltine and undergoes an extended (≥4 months) egg dormancy during late summer, autumn and winter. 4. Development of L. c. minor was observed from <1 to 15.8 °C, indicating it is a cold‐stenotherm. Its geographical distribution seems to be constrained by high summer temperatures. Within the tolerated temperature range (0–16 °C), however, temperature had no apparent effect on hatching, larval growth rate or pupal size. 5. Larval growth rate and size of pupae were significantly correlated with food availability. Completion of larval development required 300–400 degree‐days at sites where periphyton was readily available, but >800 degree‐days where periphyton availability was low. 6. No evidence for intraspecific food competition was found, whereas food competition by other blepharicerid species, mainly Hapalothrix lugubris, appeared to be a major reason for the observed food limitation. However, as L. c. minor is much more widespread than H. lugubris, food competition between these two species is likely to occur infrequently. We therefore contend that variations in the life history of L. c. minor are mainly a consequence of its thermal constraints.