Orientation of Fall-migrating Butterflies in North Peninsular Florida and Source Areas

Phoebis sennae (Pieridae), Agraulis vanillae (Heliconidae) and Urbanus proteus (Hesperiidae) migrate into peninsular Florida each fall from summer breeding areas throughout the south-eastern United States. Fall flight directions of the three species were studied at 78 sites for clues to their means of orientation and migratory routes. Mean flight directions for a visit to a site were calculated using conventional circular statistics. The distribution of flight directions permitted using linear procedures to combine visit means. The overall mean direction (OMD) of a species at a site was defined as the mean of visit means, weighted by frequency/angular variance, with a lower limit of 0.2 imposed on angular variance. In intensive studies at two sites at Gainesville, Florida, mean flight directions during fall migration were largely independent of time of day and crosswinds. The OMDs of the three species at the two Gainesville sites were similar (142 to 156°) and not significantly different among species at a site or, for the same group of dates, between sites for a species. However, day-to-day variation among species and between sites was not concordant. Visits to 65 sites on a grid throughout the southeastern United States and to 11 sites along transects inland from the Gulf and Atlantic coasts yielded 10 cases of a P. sennae OMD at a grid or transect site differing significantly from its OMD at the main Gainesville site. In each of these, the OMD was more easterly at sites inland from the Gulf coast or more southerly or south-westerly at sites inland from the Atlantic coast. By unknown means and with large, unexplained day-to-day variations in directions, P. sennae converge on peninsular Florida from widespread summer breeding grounds. They do this without closely following the Atlantic or Gulf coasts.     

Determinants of foraging profitability in two nectarivorous butterflies

ABSTRACT.

• 1 I studied flower selection and foraging energetics of Agraulis vanillae L. (Nymphalidae) and Phoebis sennae (Pieridae), two butterfly species common to north central Florida. I identified the major nectar resources exploited by several populations of these butterflies and, for each plant species, measured available nectar volumes and concentrations, corolla lengths, and density. I quantified foraging behaviour of each butterfly species at each nectar source (flower visitation rate and percentage of foraging time in flight), and used these data to estimate the net rate of energy intake of each butterfly species at each nectar source.
• 2 Estimated mean energy contents of individual flowers of the eleven exploited plant species spanned three orders of magnitude, ranging between 0.015 and 9.27 joules. Mean energy content of individual flowers was strongly correlated with mean foraging profit of both butterfly species.
• 3 Mean nectar volume strongly influenced energy content and varied widely within and among species, ranging from 0.0076 to 1.853 μ1. Nectar concentration varied between 17.1% and 40.4% sucrose-equivalents. Nectar volume was the best single predictor of foraging profitability (correlation coefficients of 0.994 and 0.984 for Phoebis and Agraulis respectively). Corolla length also strongly affected foraging profitability for both butterfly species; flower species with longer corollas were generally more profitable.
• 4 Flower density and nectar concentration showed weak or nonsignificant associations with foraging profitability.
• 5 The usefulness and limitations of these floral characteristics as bases for foraging selectivity, and the selective pressures foraging butterflies might place on the visited plants are discussed.

     

10. PROCEEDINGS OF THE SEVENTEENTH ANNUAL GENERAL MEETING OF THE S.A. ORNITHOLOGICAL SOCIETY HELD IN THE COMMITTEE ROOM OF THE PUBLIC LIBRARY. JOHANNESBURG,AT 3 P.M., SATURDAY, 8TH MARCH. 1947

Bruderer, B. 1994. Nocturnal bud migration in the Negev (Israel) a tracking radar study. Ostrich 65: 204–212.

The present publication summarizes the methodological possibilities of tracking radar and describes some features of nocturnal migration at two sites in the Negev, which include anwers to basic questions of bird migration. The directions of spring and autumn migration were practically opposite; only the headings in spring indicated some more compensation for stronger westerly winds. The volume of nocturnal spring migration was only about 65% of autumn migration, which may be an indication of mortality outside tie breeding area. Highest densities of migration at the two radar sites in the Negev Highlands (450 m above sea level) and in the Arava Valley (150 m below sea level) indicated flightlevels adjusted to atmospheric conditions aloft, and not to round level. Due to the trade-wind system, the birds heading southward in autumn flew mainly below flew mainly above 1500 m above sea level, while in spring they tended to make use of the anti-trades at higher altitudes. The decisive factor for altitude choice was the speed of tailwind in spring and autumn; other factors, such as temperature, humidity and pressure had no significant influence on the altitude distributions. With respect to the question of non-stop or intermittent flight across large desert areas, the data show that between the eastern deserts of Egypt and the Sinai/Negev complex the nocturnal migrants maintained their schedule of nocturnal flight and diurnal rest. A few exceptions of nocturnal migrants continuing migration at high altitudes into the day were identified mainly as heron- and gull-type birds. The proportion of waders and waterfowl identified by wing-beat pattern in nocturnal migration is nearly the same at both sites, indicating broad-front migration across the desert. The numbers of birds with continuous wingbeats is, however, so large compared to available estimates of waders and waterfowl wintering in Africa that careful reconsideration of the underlying assumptions in the radar and field estimates is necessary.     

Interaction between flight,reproductive development and oviposition in the pine weevil Hylobius abietis

• 1 The development of reproductive and flight capacity of pine weevils Hylobius abietis during the spring and their dispersal to, and subsequent development at, new clearfell oviposition sites comprise key phases in their life cycle in managed forests. At an old clearfell site where autumn‐emerging weevils had overwintered, weevils were trapped as they re‐emerged in the spring and tested for their ability to fly and then dissected to determine the degree of wing muscle and egg development.
• 2 Re‐emerging weevils were most abundant in pine growing at the edge of the clearfell and, over most of the trapping period (April to June), their capacity for flight (proportion flying and wing muscle width) was more advanced than in weevils from the clearfell itself, with a similar trend in the degree of reproductive development (proportion with mature eggs and egg volume).
• 3 In weevils from the clearfell, flight capacity and reproductive development increased concurrently to a peak around mid‐May. In weevils from pine, wing muscles were already well developed at the start of trapping, although few of them flew. Their more advanced development was attributed to the increased opportunities for maturation feeding after emergence in the previous autumn.
• 4 In the spring, weevils reached the canopy of trees for maturation feeding by walking and, to a lesser extent, by flight. Weevils dispersed by flight to oviposition sites in mid‐May when most of them were reproductively mature. After arrival, flight ability and wing muscle size declined rapidly but egg production was maintained until most weevils had stopped flying. When wing muscles reached their minimum size, there was a marked decline in egg size, suggesting that wing muscle breakdown is important in maintaining egg production at oviposition sites. Prospects for further wing muscle and reproductive development are discussed.

     

Autumn southward ‘return’ migration of the mosquito Culex tritaeniorhynchus in China

Abstract. Direct evidence for a southward ‘return’ migration in autumn of the mosquito Culex tritaeniorhynchus in China was obtained by aerial sampling with a net suspended from a balloon. In a preliminary study between 2 and 20 September 1990 at Jiangpu, near Nanjing, in Jiangsu Province, 11 females and 4 males of this species were taken at heights of 150–250 m. During 17–26 October 1991 at Dongxiang in northern Jiangxi Province, 44 mosquitoes (8 males, 36 females) were caught in the aerial net at heights of 80–380m in northeasterly winds (E-NNW). Most of the specimens were flying within the subsiding air-mass behind a cold front. Cx tritaeniorhynchus was the only species identified (31 females) among the mosquitoes from Dongxiang. Of 24 females dissected, 17 had N stage ovaries — interpreted as diapause, five had stage I ovaries, one had stage II ovaries, and one was gravid (stage V), but none was freshly bloodfed. Cx tritaeniorhynchus is the main vector of Japanese viral encephalitis in China, and it is possible that the virus is reintroduced to northern temperate areas in spring by northward migration of infected Cx tritaeniorhynchus females.     

Age structure and age‐related differences in molt status and fuel deposition of Dunlins during the nonbreeding season at Chongming Dongtan in east China

ABSTRACT Although most shorebirds exhibit deferred migration and deferred breeding during their first summer, Dunlins (Calidris alpina) migrate to breeding areas and breed during their first summer. First‐year and adult Dunlins should, therefore, have similar fueling and molt patterns if energetic and physiological constraints are responsible for deferred migration. From 2006 to 2008, we examined the age structure of Dunlins during the nonbreeding season at Chongming Dongtan, an estuarine wetland in the Yangtze River estuary in east China, and examined the effects of date, age, and molt status on fuel deposition during migration and during the winter. The Dunlin population at Chongming Dongtan was composed primarily of first‐year birds. Most adults and first‐year birds arrived together in late August. Regression analyses indicated that age, date, and molt status affected fuel deposition (as indicated by body mass) of Dunlins. Adults had significantly greater fuel deposits than first‐year Dunlins near the end of northward migration (May: adults 70.8 ± 6.4 g, first‐year 63.8 ± 8.0 g) and at the start of southward migration (September: adults 50.2 ± 6.1 g, first‐year 47.2 ± 4.9 g). Adults also had significantly higher fuel deposition rates than first‐year Dunlins during northward migration. Nonetheless, first‐year Dunlins migrate and breed in their first summer. Thus, other factors, such as migration distance and body size, may be more important in determining if first‐year shorebirds defer migration during their first spring and summer. During boreal spring and autumn, first‐year Dunlins in active body molt had greater body mass than those that had not initiated body molt or those in suspended molt, and premigratory fuel deposits for northward migration were greatest after prealternate molt was completed. These results suggest that body molt requires additional fuel deposits and imposes a constraint on fuel deposition for migratory flights.     

A new experimental approach to investigate migration in Pieris brassicae L.

Abstract.

• 1 It is possible to correlate distinct sequences of flight behaviour in caged females of the migrant butterfly Pieris brassicae with the main flight direction of released individuals of the same stock. The attempts to escape out of the cage point to the same mean direction as the flights of the released butterflies. This provides an experimental tool to investigate some unsolved questions concerning migration behaviour.
• 2 This study examines the constancy of the flight direction of a group of butterflies, the specificity of the direction in two populations of different geographical origin, and the frequently reported change in flight direction of subsequent generations.
• 3 The results indicate that the main flight direction of a tested group is very constant during 2 days of observation. Different directions have been detected in populations originating from Northern Germany and Southern France, respectively. The preferred directions are discussed as an adaptation to the geographical circumstances. Subsequent generations of the population from Northern Germany also show different directions. The spring generation flies to the north, the autumn generation in the opposite direction. This result corresponds to field observations that in P.brassicae a return flight occurs in subsequent generations of a year.

     

Nectar uptake rates and optimal nectar concentrations of two butterfly species

Summary The relationship between sucrose concentration of nectar and volume uptake rate by the butterflies Agraulis vanillae (Nymphalidae) and Phoebis sennae (Pieridae) was examined. Recent theoretical models simulating feeding energetics of nectarivores have assumed that this volume uptake rate is produced by a constant but undetermined pressure drop (the difference between pressure at the proximal and distal ends of the feeding channel) at all nectar concentrations. These models predict that nectar of 20–25% sucrose maximizes the rate of energy intake and should thus be preferred by nectarivores. Data collected for Agraulis and Phoebis falsify this pressure drop assumption; both species produce greater pressure drops with increasing nectar concentration. In addition, males of both species produce greater suction pressure and uptake rates than females. This results in greater rates of energy intake for males of both species. The volume uptake rates produced by each species differ from those predicted by the models. This produces a maximal rate of energy intake at 35–40% sucrose rather than 20–25%. The empirically determined relationship between energy intake rate and nectar concentration esembles that predicted for discontinuous nectar feeders such as hummingbirds more closely than the relationship predicted for continuous suction feeders, suggesting that other basic assumptions about the feeding mechanism of butterflies should be critically examined.     

Flyway‐scale analysis reveals that the timing of migration in wading birds is becoming later

1. Understanding the implications of climate change for migratory animals is paramount for establishing how best to conserve them. A large body of evidence suggests that birds are migrating earlier in response to rising temperatures, but many studies focus on single populations of model species.
2. Migratory patterns at large spatial scales may differ from those occurring in single populations, for example because of individuals dispersing outside of study areas. Furthermore, understanding phenological trends across species is vital because we need a holistic understanding of how climate change affects wildlife, especially as rates of temperature change vary globally.
3. The life cycles of migratory wading birds cover vast latitudinal gradients, making them particularly susceptible to climate change and, therefore, ideal model organisms for understanding its effects. Here, we implement a novel application of changepoint detection analysis to investigate changes in the timing of migration in waders at a flyway scale using a thirteen‐year citizen science dataset (eBird) and determine the influence of changes in weather conditions on large‐scale migratory patterns.
4. In contrast to most previous research, our results suggest that migration is getting later in both spring and autumn. We show that rates of change were faster in spring than autumn in both the Afro‐Palearctic and Nearctic flyways, but that weather conditions in autumn, not in spring, predicted temporal changes in the corresponding season. Birds migrated earlier in autumn when temperatures increased rapidly, and later with increasing headwinds.
5. One possible explanation for our results is that migration is becoming later due to northward range shifts, which means that a higher proportion of birds travel greater distances and therefore take longer to reach their destinations. Our findings underline the importance of considering spatial scale when investigating changes in the phenology of migratory bird species.

     

The long-distance migration of Nilaparvata lugens (Stål) (Delphacidae) in China: radar observations of mass return flight in the autumn

Abstract.

• 1 A case study is presented of the autumn migration of the brown planthopper, Nilaparvata lugens (Stål), in the area of Nanjing in the People's Republic of China. The study was made using a high frequency (8 mm wavelength) radar and a net suspended from a kytoon.
• 2 The observations confirmed that long-distance return migrations occur in China in mid and late September, with N.lugens being carried on the prevailing north-easterly wind towards the autumn infestation and overwintering areas of the species.
• 3 After mass take-off in the late afternoon or at dusk, the migrants flew for several hours during the evening, often in a dense layer which formed at heights between about 400 and 1000m above ground. These layers often had well-defined ceilings corresponding to an air temperature of about 16°C. The migration height was above the top of the surface temperature inversion, i.e. the migrants did not fly at the height of the warmest air.
• 4 The dense layer concentrations overflying the radar were backtracked to source areas up to 240 km away in the north-east of Jiangsu Province. Planthoppers observed emigrating from the Nanjing area would reach areas in south Anhui Province or north Jiangxi Province if they flew for 12 h.
• 5 There was a second period of mass take-off at dawn. Insect layers sometimes formed but did not last longer than 1–2h.
• 6 The present results were strikingly different from those previously observed in the dry season in the Philippines, where migratory flight durations were largely confined to periods of about 30min at dusk and dawn.
• 7 Our observations are discussed in relation to the equator-wards return migrations undertaken in autumn by other insect species, and the importance of these migrations for the maintenance of long-flying genotypes in the overwintering populations is considered.

     

Season-specific directional movement in migratory Australian butterflies

Large numbers of adults of certain species of butterfly flying in an apparently 'purposeful' manner are often noted by entomologists and the general public. Occasionally, these are recorded in the literature. Using these records we summarise information regarding the direction of movement in Australian butterflies and test whether there are consistent patterns that could account for known seasonal shifts in geographical range. The data were analysed using contingency tables and directionality statistics. Vanessa itea, Vanessa kershawi, Danaus plexippus , Danaus chrysippus and Badamia exclamationis flew predominately south in the spring–summer and north in the autumn–winter. Tirumala hamata has a strong southern component to its flight in spring but, as in Euploea core, appears non-directional in the autumn. For many supposedly known migratory species, the number of literature records are few, particularly in one season (mainly autumn). Thus, for Appias paulina , four of seven records were south in the spring–summer, as were six of nine records for Catopsilia pomona, and three of five for Zizina labradus. For Belenois java , flight records were only available for the spring and these showed geographical differences; predominantly north-west in northern Australia (Queensland) and south-west in southern Australia (Victoria, New South Wales). There were too few records for Papilio demoleus in the literature (four only) to draw any conclusions. Major exceptions to the seasonal trend of south in the spring and north in the autumn were Junonia villida , which showed a predominant north-westward direction in both seasons, and Eurema smilax, with a predominant southern or western flight in both seasons. We discuss these species specific trends in migration direction in relation to seasonal shifts in suitable habitat conditions, possible cues used in orientation and in timing changes in direction.     

Flower constancy and learning in foraging preferences of the green-veined white butterfly Pleris napi

Abstract.

• 1 Evolutionary pressure should select for efficient foraging strategies, within the constraints of other selective forces. We assess the mechanisms underlying flower choice in the butterfly, Pieris napi (L.), which as an adult forages for nectar. Experiments were carried out on a laboratory colony, using artificial flowers of two colours, and replicated on two successive generations.
• 2 When nectar was freely available from all flowers, equal numbers of butterflies visited each colour, but individual butterflies exhibited flower constancy, showing a strong preference for one colour or the other.
• 3 Following 3 day conditioning periods in which nectar was available from flowers of one colour only, butterflies responded by developing a preference for this colour, which persisted when both flower colours were refilled. This preference could subsequently be switched to the other flower colour following a further 3 days of conditioning. These are interpreted as adaptive (learned) responses, which would have obvious selective benefits in the field, enabling butterflies to avoid flower species which experience has shown are poor sources of nectar, and to adapt to temporal and spatial changes in nectar availability.

     

Seasonal migration strategies of Common Ringed Plovers Charadrius hiaticula

How individual birds schedule their movements and use different sites during the non‐breeding season are fundamental issues in avian migration ecology, and studies have often revealed strong seasonal variation in such strategies. Using geolocators we tracked Common Ringed Plovers Charadrius hiaticula from northern Norway to West Africa and back to assess whether there were differences in migratory speed, duration and stopover use between autumn and spring migration and whether birds used multiple sites during the non‐breeding season. Although the pace of migration was similar between autumn and spring, the length of flight bouts and duration of the preceding stopovers were positively correlated only in autumn. Four of five birds showed a marked southward movement in mid‐winter.     

Breeding latitude predicts timing but not rate of spring migration in a widespread migratory bird in South America

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Spring remigration of the monarch butterfly, Danaus plexippus (Lepidoptera: Nymphalidae) in north-central Florida: estimating population parameters using mark-recapture

Monarch butterflies [Danaus plexippus) of the eastern North American population migrate each fall from die northern U.S.A. and southern Canada to overwintering sites in Mexico and return the following spring to the southeastern U.S.A. where they lay eggs and then die. The spring remigration is the least studied phase in the annual migration cycle. We therefore conducted a mark-recapture study and examined population recolonization dynamics and residence time in a north-central Florida pasture where the monarch's milkweed host plant (Asclepias humistrata) was abundant. Beginning in late March 1995 two waves of monarchs arrived, their numbers peaked at 71 individuals by mid-April, and the butterflies disappeared in early May. After arriving, the adults remained for 3–5 days, laid eggs and then continued to migrate. We also compared population sizes and arrival times in 1994 and 1996. We found no evidence of a second spring generation, which was also consistent with the deteriorating quality of the A. humistrata plants. Individuals of the new spring generation disappear shordy after eclosion. The arriving population was approximately nine times greater in 1995 than in 1996. Our findings support two recent hypodieses: (1) the bird-like migration of the monarch butterfly in North America evolved with the northward expansion and phenology of milkweeds; and (2) monarchs appear to be migratory throughout their annual cycle of several generations. By lingering for only a short time at each milkweed patch they encounter, the old monarchs returning from Mexico locate the resurgent milkweed flora over an extensive area in the southern states. Then, within less than a month, their fresh offspring continue the migration and exploit the unfolding cornucopia of milkweeds as the spring advances northward. The more we discover about the biology of this insect, the more remarkable is its annual migratory, breeding and overwintering cycle.     

Adverse wind conditions during northward Sahara crossings increase the in‐flight mortality of Black‐tailed Godwits

Long‐distance migratory flights are predicted to be associated with higher mortality rates when individuals encounter adverse weather conditions. However, directly connecting environmental conditions experienced in‐flight with the survival of migrants has proven difficult. We studied how the in‐flight mortality of 53 satellite‐tagged Black‐tailed Godwits (Limosa limosa limosa) during 132 crossings of the Sahara Desert, a major geographical barrier along their migration route between The Netherlands and sub‐Saharan Africa, is correlated with the experienced wind conditions and departure date during both southward and northward migration. We show that godwits experienced higher wind assistance during southward crossings, which seems to reflect local prevailing trade winds. Critically, we found that fatal northward crossings (15 deaths during 61 crossings) were associated with adverse wind conditions. Wind conditions during migration can thus directly influence vital rates. Changing wind conditions associated with global change may thus profoundly influence the costs of long‐distance migration in the future.     

The migration of the great snipe Gallinago media: intriguing variations on a grand theme

The migration of the great snipe Gallinago media was previously poorly known. Three tracks in 2010 suggested a remarkable migratory behaviour including long and fast overland non‐stop flights. Here we present the migration pattern of Swedish male great snipes, based on 19 individuals tracked by light‐level geolocators in four different years. About half of the birds made stopover(s) in northern Europe in early autumn. They left the breeding area 15 d earlier than those which flew directly to sub‐Sahara, suggesting two distinct autumn migration strategies. The autumn trans‐Sahara flights were on average 5500 km long, lasted 64 h, and were flown at ground speeds of 25 m s^?1 (90 km h^?1). The arrival in the Sahel zone of west Africa coincided with the wet season there, and the birds stayed for on average three weeks. The birds arrived at their wintering grounds around the lower stretches of the Congo River in late September and stayed for seven months. In spring the great snipes made trans‐Sahara flights of similar length and speed as in autumn, but the remaining migration through eastern Europe was notably slow. All birds returned to the breeding grounds within one week around mid‐May. The annual cycle was characterized by relaxed temporal synchronization between individuals during the autumn–winter period, with maximum variation at the arrival in the wintering area. Synchronization increased in spring, with minimum time variation at arrival in the breeding area. This suggests that arrival date in the breeding area is under strong stabilizing selection, while there is room for more flexibility in autumn and arrival to the wintering area. The details of the fast non‐stop flights remain to be elucidated, but the identification of the main stopover and wintering areas is important for future conservation work on this red‐listed bird species.     

Routes of migrating soaring birds

Soaring migrants travelling through Israel use three principal routes which are used in the opposite directions during the spring and autumn: (1) the Western Route lies mainly along the western edge of the central mountain range, (2) the Eastern Route lies mainly along the Jordan Valley, crossing the mountain range during part of the day, continuing southward along the Dead Sea towards the Sinai, and joining the Western Route in autumn and (3) the Southern-Elat Mountains Route. The geomorphological structure of Israel, with a central mountain range dividing the country roughly into three landscape units, plays a central role in route selection. In the autumn, the Western Route migration axis is deflected at the beginning of the day from east to west for 10–25 km, depending on weather conditions and the flock's roosting locations. Between 10.00 h and 11.00 h, the daily breeze blowing from the Mediterranean Sea influences the migration axis, which is slowly deflected back to the east. A parallel deflection of the migration axis occurs in the Eastern Route in the autumn. The route moves southwest over the eastern slopes of the central mountain range during the morning hours and over the slope, which absorbs direct radiation from the sun, creating good soaring conditions. Towards late afternoon, when the breeze from the sea starts, the axis is deflected to the east, to the Jordan Valley. In the Elat Mountains, the wind flow plays a similar role, but because the topography of the southern Arava Valley causes a change in wind direction, the axis moves during the day in a north-south direction. In addition to the axis movement on a daily scale, a seasonal deflection of the migration axis from east to west also exists. During autumn migration, early migrants (e.g. White Storks Ciconia ciconia) tend to travel on an eastern route, while late migrants (e.g. White Pelican Pelecanus onocrotalus) travel along the Mediterranean coast. This fluctuation was probably because of sub-optimal soaring conditions along the coastal plain during August. In September, temperature differences between the sea and land decrease and the influence of the marine inversion gradually declines, until its influence disappears completely in October. A comparison of the numbers of soaring birds seen over Israel in the autumn and spring shows significant seasonal differences in the use of the various routes. For example, only one species, the Steppe Eagle Aquila nipalensis, flies over the Elat Mountains in the autumn, compared to more than 30 species in the spring. In the autumn, White Storks pass over only along the Jordan Valley axis, whereas in the spring, about half the migrating storks also pass over the western edge of the central mountain range. Honey Buzzards Pernis apivorus fly along the Western Route in large numbers in the autumn, while concentrating almost totally over the Elat Mountains in the spring. These differences are related to the global migration routes between the breeding and the wintering grounds in relation to the Red Sea, which birds avoid crossing, thus causing them to follow different routes in autumn, and spring.     

Temporal changes in host adaptation in the pea aphid, Acyrthosiphon pisum

Abstract.

• 1 Temporal changes in host adaptation were followed in a local population of the pea aphid, Acyrthosiphon pisum. Aphid clones were collected in one alfalfa and one clover field at three different times. In the spring, first-generation females were collected. Later, in the autumn, females belonging to the last parthenogenetic generation were collected. Lastly, sexual females were collected after mating in autumn and allowed to produce eggs which were hatched. The performance was evaluated on alfalfa and clover. The spring-collected individuals were also assessed on peas.
• 2 On the overwintering hosts clover and alfalfa, the clones performed best on the plant of origin, i.e. negative correlations in performance. Correlations between performance on the temporary summer host, pea, and that on clover/alfalfa were weak or nonsignificant.
• 3 Significant variation in host performance was found within both host fields at spring, which is a prerequisite for changes in clone composition due to selection/migration.
• 4 The clones from alfalfa showed an increase in mean performance on alfalfa between spring and autumn, whereas no changes among the clones from the clover field were observed. This difference in seasonal response between the two fields could have been the result of larger variation in performance among the alfalfa clones and/or a differential tendency to migrate among clones in both fields.
• 5 After sexual recombination in the autumn, mean performance in the alfalfa field returned to the spring level, probably as a result of emergence of new genetic combinations. In the clover field, mean performance did not change significantly over time.