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Migratory animals exhibit traits that allow them to exploit seasonally variable habitats. In environments where migration is no longer beneficial, such as oceanic islands, migration-association traits may be selected against or be under relaxed selection. Monarch butterflies are best known for their continent-scale migration in North America but have repeatedly become established as nonmigrants in the tropical Americas and on Atlantic and Pacific Islands. These replicated nonmigratory populations provide natural laboratories for understanding the rate of evolution of migration-associated traits. We measured >6,000 museum specimens of monarch butterflies collected from 1856 to the present as well as contemporary wild-caught monarchs from around the world. We determined 1) how wing morphology varies across the monarch's global range, 2) whether initial long-distance founders were particularly suited for migration, and 3) whether recently established nonmigrants show evidence for contemporary phenotypic evolution. We further reared >1,000 monarchs from six populations around the world under controlled conditions and measured migration-associated traits. Historical specimens show that 1) initial founders are well suited for long-distance movement and 2) loss of seasonal migration is associated with reductions in forewing size and elongation. Monarch butterflies raised in a common garden from four derived nonmigratory populations exhibit genetically based reductions in forewing size, consistent with a previous study. Our findings provide a compelling example of how migration-associated traits may be favored during the early stages of range expansion, and also the rate of reductions in those same traits upon loss of migration.

Monarch butterfly wing morphology varies substantially throughout their global range, both between resident and migratory populations and also within the migratory North American population. Here, we use a dataset comprising more than 1800 North American individuals collected between 1878-2017 to characterize the factors shaping continent-wide patterns of wing morphological variation. North American overwintering butterflies have forewings that are approximately 4.4% larger than those collected in summer breeding areas. Monarchs overwintering in Mexico have forewings that are approximately 1.8% larger than monarchs overwintering in California, conducive to the idea that migration distance is positively correlated with wing area. We find evidence for a latitudinal cline within North America, such that butterflies collected at higher latitudes have significantly larger and more elongated forewings. We also find a significant increase of approximately 4.9% in forewing area between 1878-2017, but no difference through time in wing elongation. This result is corroborated by a reanalysis of a recently published dataset of more than 600 butterflies from Mexican overwintering sites. We discuss possible reasons for this increase in wing size through time, including northward shifts in the monarch’s breeding range and changes in relative abundance of milkweed hosts, and present experimental data addressing the influence of larval host plant on adult wing morphology. Our analysis suggests that (1) migration is indeed an important selective force for monarch wing morphology; (2) wing size has increased through time in North America; (3) factors such as host plant identity must be considered to fully understand monarch wing morphological variation.

To broaden perspectives and stimulate research on migration, I survey the bird species that breed in the northern hemisphere and migrate to the southern hemisphere and species that migrate within the southern hemisphere, comparing routes, seasonal patterns and life histories. Differences in the area and latitudinal extent of land masses on the two sides of the Equator influence patterns of bird migration. In contrast to birds breeding in the northern hemisphere, no land or freshwater birds breeding in the southern hemisphere migrate between continents and only a very few cross the Equator. Furthermore, except for shorebirds, few northern intercontinental migrants reach the southern hemisphere in regions south of the equatorial forest belt, because most encounter, and are filtered out by, suitable habitats en route. Australasia is an extreme case because only 10 land or freshwater migrants from the northern hemisphere regularly occur there (most are uncommon or rare) compared with 42 in Africa and 28 in South America, and no Australasian breeders enter Asia beyond Wallace’s Line. Historical geographical and oceanic barriers may be an additional factor limiting migration to Australasia. There are generally no or only slight differences in frequencies of austral migrants within foraging guilds or families across southern continents. Exceptions are rallids, with more migrants in Africa, and cuckoos and nectarivores, with more Old World than New World migrants. Austral migrations are of shorter distances than most of those of the northern hemisphere, and they appear to vary more with respect to routes and patterns. Breeding and non-breeding ranges frequently overlap. Partial migration is common, but there is no evidence that it differs in frequency from that in northern regions. Because climate is generally milder and drier in the southern hemisphere, rainfall is a more important influence on migration than in the north especially in some nomadic birds, but temperature also predicts migration frequency and pathways for many species. These patterns are similar across southern continents, but each continent has its own characteristics. Southern hemisphere migrants seem to display ecophysiologies and orientation mechanisms similar to those found in northern hemisphere species, but very few southern species have been studied. I argue that the variation present among southern hemisphere migrations provides exceptional opportunities to understand the evolution and ecology of migration systems. In order to take advantage of these systems, we need to focus on variation in movement behaviour, on associated syndromes of traits, and on the particular features of natural selection and ecology setting thresholds that lead to the diverse migration patterns observed.

We outline a general scheme for migration that applies across taxa, incorporates the several varieties of migration, and includes all levels of biological organization, from genes to populations. The scheme links the environment, pathways, traits, and genes, and highlights the selective forces that shape and maintain migratory adaptation. We endorse an individual-based behavioral definition of migration that allows an objective distinction between migration and other forms of movement. We recognize migration as an adaptation to resources that fluctuate spatiotemporally either seasonally or less predictably, and note that it is often preemptive. Migration plays a central role in the spatial dynamics of mobile populations, and is largely distinct in both form and function from the within-population mixing arising from postnatal dispersal and from the interpatch movements characteristic of metapopulations. We call for more interaction between biologists studying different taxa and different forms of movement, and between behaviorists and population ecologists.

Evolutionary change results from selection acting on genetic variation. For migration to be successful, many different aspects of an animal’s physiology and behaviour need to function in a co-coordinated way. Changes in one migratory trait are therefore likely to be accompanied by changes in other migratory and life-history traits. At present, we have some knowledge of the pressures that operate at the various stages of migration, but we know very little about the extent of genetic variation in various aspects of the migratory syndrome. As a consequence, our ability to predict which species is capable of what kind of evolutionary change, and at which rate, is limited. Here, we review how our evolutionary understanding of migration may benefit from taking a quantitative-genetic approach and present a framework for studying the causes of phenotypic variation. We review past research, that has mainly studied single migratory traits in captive birds, and discuss how this work could be extended to study genetic variation in the wild and to account for genetic correlations and correlated selection. In the future, reaction-norm approaches may become very important, as they allow the study of genetic and environmental effects on phenotypic expression within a single framework, as well as of their interactions. We advocate making more use of repeated measurements on single individuals to study the causes of among-individual variation in the wild, as they are easier to obtain than data on relatives and can provide valuable information for identifying and selecting traits. This approach will be particularly informative if it involves systematic testing of individuals under different environmental conditions. We propose extending this research agenda by using optimality models to predict levels of variation and covariation among traits and constraints. This may help us to select traits in which we might expect genetic variation, and to identify the most informative environmental axes. We also recommend an expansion of the passerine model, as this model does not apply to birds, like geese, where cultural transmission of spatio-temporal information is an important determinant of migration patterns and their variation. [KEYWORDS: Individual variation ; Migration ; Migratory syndrome ; Migratory traits ; Quantitative genetics]

Ornithologists, and especially northern hemisphere ornithologists, have traditionally thought of migration as an annual return movement of populations between regular breeding and non-breeding grounds. Problems arise because selection does not ordinarily act on populations and because organisms of many taxa (including birds) are clearly migrants, but fail to undertake movements of the kind described. There are also extensive return movements that are not migratory. I propose that it is more useful to think of migration as a syndrome of behavioral and other traits that function together within individuals, and that such a syndrome provides a common ground across taxa from aphids to albatrosses. Large-scale return movements of populations are one outcome of the syndrome. Similar behavioral and physiological traits serve both to define migration and to provide a test for it. I use two insect (Hemipteran) examples to illustrate migratory syndromes and to demonstrate that, in many migrants, behavior and physiology correlate with life history and morphological traits to form syndromes at two levels. I then compare the two Hemipterans with migration in birds, butterflies, and fish to assess the question of whether there are migratory syndromes in common between these diverse migrants. Syndromes are more similar at the level of behavior than when morphology and life history traits are included. Recognizing syndromes leads to important evolutionary questions concerning migration strategies, trade-offs, the maintenance of genetic variance and the responses of migratory syndromes to both similar and different selective regimes.

Soapberry bugs are worldwide seed predators of plants in the family Sapindaceae. Australian sapinds are diverse and widespread, consisting of about 200 native trees and shrubs. This flora also includes two introduced environmental weeds, plus cultivated lychee (Litchi chinensis Sonn.), longan (Dimocarpus longan Lour.) and rambutan (Nephelium lappaceum L.). Accordingly, Australian soapberry bugs may be significant in ecology, conservation and agriculture. Here we provide the first account of their ecology. We find five species of Leptocoris Hahn in Australia, and list sapinds that do and do not serve as reproductive hosts. From museum and field records we map the continental distributions of the insects and primary hosts. Frequency of occupation varies among host species, and the number of hosts varies among the insects. In addition, differences in body size and beak length are related to host use. For example, the long-beaked Leptocoris tagalicus Burmeister is highly polyphagous in eastern rainforests, where it occurs on at least 10 native and non-native hosts. It aggregates on hosts with immature fruit and commences feeding before fruits dehisce. Most of its continental range, however, matches that of a single dryland tree, Atalaya hemiglauca F. Muell., which has comparatively unprotected seeds. The taxon includes a smaller and shorter-beaked form that is closely associated with Atalaya, and appears to be taxonomically distinct. The other widespread soapberry bug is the endemic Leptocoris mitellatus Bergroth. It too is short-beaked, and colonises hosts phenologically later than L. tagalicus, as seeds become more accessible in open capsules. Continentally its distribution is more southerly and corresponds mainly to that of Alectryon oleifolius Desf. Among all host species, the non-native environmental weeds Cardiospermum L. and Koelreuteria Laxm. are most consistently attacked, principally by L. tagalicus. These recent host shifts have biocontrol implications. In contrast, the sapinds planted as fruit crops appear to be less frequently used at present and mainly by the longer-beaked species.

The standard model for the migration of the monarch butterfly in western North America has hitherto been movement in the autumn to overwintering sites in coastal California, followed by a return inland by most individuals in the spring. This model is based largely on observational and limited tagging and recovery data. In this paper we test the model by plotting many years of museum and collection records on a monthly basis on a map of the region. Our plots suggest a movement of Oregon, Washington and other north-western populations of summer butterflies to California in the autumn, but movement of more north-easterly populations (e.g. from Idaho and Montana) along two pathways through Nevada, Utah and Arizona to Mexico. The more westerly of these two pathways may follow the Colorado River south as indicated by museum records and seasonal temperature data. The eastern pathway may enter northern Utah along the western scarp of the Wasatch Mountains and run south through Utah and Arizona. Further analysis of distributions suggests that monarch butterflies in the American West occur primarily along rivers, and there are observations indicating that autumn migrants often follow riparian corridors. More data are needed to test our new model; we suggest the nature of the data required. (c) 2005 The Linnean Society of London.

We investigated the adaptive significance of behavioural thermoregulation in univoltine populations of the grasshopper Melanoplus sanguinipes along an altitudinal gradient in California using laboratory tests of animals raised under different temperatures. Trials consisted of continuous body temperature measurements with semi-implanted microprobes in a test arena, and observation and simultaneous recording of behavioural responses. These responses included mobility, basking and orientation of the body axes (aspect angle) towards a radiation source. Mobility and basking are determined by the altitudinal origin of the parental generation and not by the temperature treatments. With increasing altitude, individuals tend increasingly to raise body temperatures via mobility and increased basking. In contrast, body orientation towards the radiation source is influenced by the temperature treatments but not by the altitude of origin. Individuals experiencing higher temperatures during rearing show a lower tendency to lateral flanking. We conclude that body orientation responses are not adapted locally. In contrast other components of the behavioural syndrome that increase body temperature, such as mobility and basking, are adaptive in response to local selection pressure. The thermoregulatory syndrome of these grasshoppers is an important contribution to life-history adaptations that appropriately match season lengths.

The colonization of an introduced plant by s bugsin Florida hasres ulted in the rapid evolution of a distinctive "host race." Differences in the synchrony and persistence of local seed crops may have favored interracial differentiation in sight/life history tactics. To advance previous work, we quantiÞed seed availability in individual native and introduced host plants. We tested the prediction that the new hostOs relatively discrete period of annual seed production has selected for a higher frequency of a short-winged, sightless morph with a briefer generation time. Contrary to that prediction, short-winged bugs were not more common in the derived race, but further investigation revealed the unexpected presence of a long-winged morph, that like the short-winged, lacks sight muscle and exhibits the same rapid life cycle advantages. Consistent with prediction, the derived race, descended from volant long-winged colonists, shows an abundance of this "cryptic" sightless morph. In total, four sight/life history morphs were discovered, including two additional long-winged types that either histolyze or retain sight muscle. The morphs differed in life history traits both within and between host races. Morph frequency may be insuenced by evolution in correlated characters: we found that beak length, which has rapidly evolved to be shorter on the smaller fruits of the introduced host, is discontinuously distributed between sight morphs, being greater in long- winged bugs. This study shows complex, unpredicted evolutionary relations between a mobility character, a trophic character, and the life history.

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.

With reciprocal rearing experiments, we tested the hypothesis that adaptive differences in host-use traits among soapberry bug populations have a genetic basis. These experiments were conducted with two host races from Florida, an ‘ancestral-type’ one on a native host plant species and a ‘derived’ one on a recently introduced plant species (colonized mainly post-1950), on whose seed crops this insect depends for growth and reproduction. Compared to the native host species, the introduced host produces larger seed crops over a much briefer annual period. Its seeds are also significantly higher in lipids and lower in nitrogen. The bug populations exhibit greater juvenile survivorship on their ‘home’ hosts; that is, the derived population survives better on seeds of the introduced host than does its ancestral-type counterpart, and vice versa. Regardless of the rearing host, populations from the introduced host lay much smaller eggs, and fecundity measures show a more complex pattern than does survivorship: the ancestral-type population produces eggs at the same rate on each host, while the derived population is less fecund on the native host and exhibits enhanced fecundity on the introduced host. These results indicate that the population differences are evolved rather than host-induced. They appear to be adaptive responses to host differences in the spatial and temporal distribution of seed availability and nutritional quality, and show that increased performance on the alien host has evolved with surprising speed and magnitude, with concomitant reductions in performance on the original host.

Insects display much variation in life histories mediated by juvenile hormone. We focus on the contribution of JH to variations in migratory life histories. In many migrants such as the large milkweed bug and the monarch butterfly, JH directly influences migratory flight and the relation between flight and reproduction (oogenesis-flight syndrome). In the true armyworm, JH regulates interactions among female calling, pheromone production, ovarian development, and migration with varying blends of structurally related forms of JH and JH acid. A role for JH also occurs in wing polymorphisms. Aphids regulate wing production via JH-mediated maternal effects; and in crickets, JH esterase modulates the JH influence on wing form. In addition, JH is implicated in wing muscle histolysis. The comprehensive Fairbairn model for JH regulation of wing polymorphisms in flight behavior predicts that JH action will depend on the mode of genetic control, whether single locus or polygenic. Our own studies of the soapberry bug, Jadera haematoloma, reveal a four-morph wing polymorphism in a species rapidly evolving on a new host plant. There are long- and short-winged forms, and the long-winged form displays three degrees of flight muscle histolysis. The polymorphism is subject to both genetic and environmental variations that are mediated by JH. Application of methoprene increases the frequency of the short-winged forms, but there is both within- and between-population genetic variation and genotype by environment interaction (plasticity) in the response to JH. Arch. Insect Biochem. Physiol. 35:359–373, 1997. © 1997 Wiley-Liss, Inc.

In this contribution we consider the biology of invading organisms after they have become established. Adaptive radiation over the long term has been a favorite subject in evolutionary biology. Examples have been Darwin's finches in the Galapagos and the honeycreepers and Droscphila of Hawaii. Hawaiian honeycreepers have evolved from a finch-like ancestor into guilds of seed eaters, nectar feeders, and combined nectar and insect feeders plus some species with unique beak structures. In the Hawaiian Drosophila sexual selection may have driven the extensive adaptive radiation and speciation in the group. The North American soapberry bug Jadera haemotoloma is an interesting model for post-invasion evolution in the short term. Some populations have moved onto introduced goldenrain trees Koelreuteria spp. and have evolved different stylet (mouthpart) lengths, as a function o f fruit size, and new host preferences, all within the last 50 years. These rapid responses are possible because of high additive genetic variances for these traits. Similarly, there has been rapid evolution of life history variation in American shad introduced from east coast to west coast rivers. We postulate that invaders most likely to integrate successfully are those in which high levels of additive genetic variation are expressed in traits most likely to be adaptive in the new environment. Copyright © 1996 Published by Elsevier Science Limited

Embryonic development times and the stage at which embryonic diapause occurs varied dramatically among 23 populations of the Melanoplus sanguinipes/ devastator species complex in California, USA. Grasshoppers were collected from a wide range of latitudes (32°57N to 41°20N) and altitudes (10m to 3031 m), spanning much of the variation in climatic conditions experienced by these insects in California. When reared in a “common garden” in the laboratory, total embryonic development times were positively correlated to the mean annual temperature of the habitat from which the grasshoppers were collected (varying from about 19 days to 32 days when reared at 27°C). These grasshoppers overwinter as diapausing eggs and the proportion of embryonic development completed prior to diapause was significantly higher in populations collected from cool habitats (>70%) than in populations collected from warm environments (

This paper applies the biological species concept to two ecologically distinct species of grasshopper, Melanoplus sanguinipes and M. devastator, by testing for reproductive isolation in the field in California. Two independent techniques for assessing gene flow between species were employed. Firstly, we examined male genitalic morphology in populations from the foothills of the Sierra Nevada where the two species are parapatric. Two genetically based genitalic traits that differed between allopatric populations of each species formed a cline in this zone. Males captured in the field from the region of parapatry resembled offspring from interspecific laboratory hybridizations. Secondly, we surveyed electrophoretic variation of populations from across California and used F statistics to estimate levels of gene flow within and between species. Results from both morphology and F statistics suggested that these grasshoppers are not reproductively isolated in nature but that gene flow between species is reduced relative to within-species. These field results were consistent with a laboratory study that showed partial but not complete hybrid egg inviability. Because the hybrid zone is centred along an ecological transition, this system offers an opportunity to investigate whether adaptive changes across the zone contribute to reduced gene flow between species.

Three primary approaches have been used to study the genetics of migration: the analyses of population differences, of single locus effects, and of polygenic influences. Studies of populations reared under similar conditions in “common garden” experiments frequently reveal gene effects contributing to differences in migratory tendency. Single locus effects are known, but are not common, a result to be expected given that migration is complex. Quantitative genetic studies reveal that heritabilities for migration related traits are often high (approximately 0.5 or more) suggesting significant amounts of genetic variation on which natural selection can act. Analyses of genetic correlations demonstrate that migratory behavior is part of a syndrome that includes aspects of both physiology and life history traits. The latter are characteristically those which contribute to colonizing ability. Migratory behavior thus does not evolve in isolation. New migration patterns are still evolving, as would be predicted from observed environmental changes and the genetic variation present in migratory species.

Natural selection can occur when phenotypic variation in a trait related to fitness is heritable (26, 39). The trait can be simple (e.g. appendage length) or complex (e.g. shape, reaction norms), but as long as individuals vary in their expression, and such variance is genetically based, natural selection can act. If over several generations natural selection is consistent and unconstrained by other factors such as antagonistic pleiotropy (145) or physiological thresholds (170), then a population will evolve. Although selection acts upon phenotypic variation, the resulting changes in genotypic frequencies affect evolution. Thus, the developmental mechanisms linking phenotype to genotype are of fundamental concern to an understand­ ing of evolutionary change. Because development is an elaborative process and adult phenotype is a summation of activities accrued over the life span of the individual, developmental events occurring early in the life cycle can significantly influence the phenotype at later stages. In some cases, such developmental variation will be dampened as a result of homeostatic mech­ anisms [e.g. targeted growth of body size in mice (136), brain size allometry (135), or developmental programming (183)], while in others developmental

Life history theory predicts that migratory fishes should delay reproduction, be larger at first reproduction, and have higher fecundities than nonmigrants. We tested this hypothesis by comparing life histories of anadromous ("estuary") and resident freshwater ("upstream") threespine sticklebacks (Gasterosteus aculeatus L.) from the Navarro River, California, USA. Using a split-brood, two-environment breeding design, families from cach population were divided and reared in both freshwater and seawater overwintering environments. In both treatments, the more migratory estuary sticklebacks were larger at first reproduction and had large initial clutch sizes; in the freshwater treatment, the estuary sticklebacks matured later than the upstream fish. Population means varied little across treatments, indicating that the average effects of the different overwintering conditions were slight. The responses of individual families to a given overwintering treatment were highly variable in both populations, as reflected in significant family x treatment effects for all traits. Phenotypic correlations among life history traits were significant and positive for most traits, and were similar in magnitude in both populations. Differences in the relative degree of specialization for migration may in part explain variation in life history between these populations.

A field study of the relationship between host plant phenology and the reproductive pattern of the large milkweed bug, Oncopeltus fasciatus, was conducted in south Florida. Since O. fasciatus need seeds of either milkweed or Nerium oleander plants to reproduce, reproduction takes place on only those host plants that are producing seed pods. Two of four major host plants, Asclepias incarnata and Sarcostemma clausa fruit seasonally, producing pods in early autumn and early winter, respectively. The third milkweed host, Asclepias curassavica, produces almost no pods midsummer (although it flowers abundantly) and few pods midwinter. Nerium oleander (Apocynaceae) produces some pods all year but is only used by O. fasciatus in the summer when milkweeds are not producing pods. Correspondingly, reproduction of O. fasciatus has been observed year round, but relatively few females reproduce in midwinter, coinciding with decreased pod production and low temperatures. This pattern is consistent with the hypothesis that a photoperiodic cue of short day lengths under conditions of cool temperatures may cause adult females to enter diapause and delay reproduction in the field. A comparison of plant phenologies and rainfall between 1976, a very dry year, and 1978, a year with normal rainfall, showed that extreme dryness disrupted the seasonal fruiting of the milkweeds and consequently the reproduction of O. fasciatus.

Daily cyclic activities in various behaviours were similar in three species of tropical Oncopeltus (Heteroptera, Lygaeidae). Opposition in females and general movement in both sexes peaked in the middle to late light period. In contrast, feeding and mating peaked at the end of the light period and the beginning of the dark respectively. Other behaviours also tended to display one or the other of these patterns. The diel timing of behaviour observed in this study essentially duplicated the periodicities of a migratory population of Oncopeltus fasciatus from Iowa. The similarity of results across species and populations suggests a general pattern for the genus which probably serves to synchronize behaviours with environmental cycles and appropriate physiology. This similarity is interesting in view of the broad geographical range of the genus across both temperate zone and tropics and in view of the differences among populations with respect to photoperiodic determination of diapause.

Differences were found in duration of diapause, as measured by days to first oviposition following adult eclosion, between female Oncopeltus fasciatus (Dallas) mated with male O.sandarachatus (Say) (mean =55 days), and those mated with their brothers (mean =99 days). Under the same conditions (LD 11:13 at 23°C), O.sandarachatus pairs showed no delay in reproduction (mean = 12 days). Male O.sandarachatus exhibit greater sexual activity than male O.fasciatus, and it is postulated that this behaviour may play a significant role in determining the length of diapause in the female.

The plant and insect communities of early, secondary succession beginning with bare ground in an Old World site (southern Britain) and a New World site (Iowa, U.S.A.) shared a number of characteristics. Both sites showed similar temporal patterns of plant species cover and species richness, although overall richness was greater at the Old World site. Annuals dominated at both sites during the first year of succession and were largely replaced by perennials in the second year. Monocotyledons were more abundant at the Old World site, especially in the second year. The two sites differed markedly in the contribution of native and introduced plant species, with the Old World site dominated by natives and the New World site by alien plant species. Insect herbivore load was greater at the Old World site, when expressed in terms of structural complexity of the vegetation, suggesting that there may be major differences in the influence of herbivores on the direction and rate of succession at the two sites.

Using two geographically adjacent, temperate-zone populations of the milkweed bug Oncopeltus fasciatus, we employed a split-family experimental design to estimate genotype-environment interaction (g-e) and across-environment genetic correlations for two female life history characters, age at first reproduction and fecundity over the first 5 days of reproduction. For each of a number of families, some sisters were reared in a short-day environment (LD 11:13, 23$^\circ$C) and others in a long-day environment (LD 14:10, 23$^\circ$C); the environments were chosen to simulate the temporal environmental heterogeneity experienced by lineages of temperate-zone bugs. Bugs normally exhibit a plastic response to such heterogeneity by delaying the onset of reproduction in short days relative to that in long days, the delay providing time for migration to occur. In one population, there is significant variation caused by g-e for the age at first reproduction, indicating the existence of genetic variance for plasticity...

This study was designed to examine life history flexibility arising from phenotypic plasticity in response to temperature and from maternal effects in response to reproductive diapause in a temperate zone population of the milkweed bug (Oncopeltus fasciatus). We employed a split-family, first-cousin, full-sib design with siblings reared at different temperatures in order to quantify phenotypic plasticity, maternal effects, and variation for each. The following traits were analyzed: development time, age at first reproduction, longevity, early-life fecundity, and wing length. We found both life history plasticity and maternal effects on life history traits which tend to enhance the colonizing ability of offspring born to mothers that have undergone reproductive diapause. We were unable to demonstrate additive genetic variation for plasticity for any of the traits, while for development time and wing length we found variation due to non-additive genetic or common-environmental sources. We were also unable to demonstrate additive genetic variation for maternal effects, although variation may exist at low levels that are difficult to detect using cousin-families. The apparent lack of variation in this population would constrain evolution of life history flexibility even though considerable flexibility exists in the phenotype.

By eliminating the food plant, Asclepias curassavica, monarch butterflies, Danaus plexippus, have virtually eliminated milkweed bugs, Oncopeltus spp., from the island of Barbados. The relatively open terrain of Barbados means the plants have no refuge; the butterflies survive on an alternate milkweed food plant, Calotropis procera, whose thick-walled pods make seeds unavailable to the bugs.

Complete sets of life-history data (sufficient to construct life-tables and calculate intrinsic rates of increase) were collected at each of three constant temperatures for descendants of two tropical populations of the Large Milkweed Bug, Oncopeltus fasciatus. Although the two populations occur only about 60 km apart, they experience quite different thermal regimes, with little variation in mean monthly temperature at either site. In addition to the pronounced effect of ambient temperature on life-history traits, significant population-by-temperature interactions were observed for six of the eight traits examined. The data and the recent history of the species' distribution are consistent with the hypothesis that natural selection in the cool habitat has favored improved survival and increased reproduction at cool temperatures, with some trade-offs with respect to performance at higher (but ecologically relevant) temperatures.

1. Unilateral removal of the eyestalk (optic ganglia and medulla terminalis) in the Bermuda spiny lobster, Panulirus argus, disrupts normal initiation of feeding activity via chemo-tactile stimulation of the antennule on the side of the ablation. This deficit may be permanent for it has lasted without apparent remission for over five months and two molts. 2. Unilateral eyestalk ablation also produces a temporary increase in antennular cleaning activity directed toward the antennule on the side of the ablation. This effect seems to last for less than 16 days. 3. Unilateral eyestalk ablation does not appreciably disturb spontaneous antennular movements or responses to mechanical stimulation of the antennule on either side. Nor does it destroy the ability of the lobster to give differential responses to mechanical and chemo-tactile stimuli. 4. Most lobsters recover normal sensory and motor functions in antennules that regenerate after amputation of the distal segment and sensory flagellae. In about 1/3 of the animals, however, some form of sensory or motor abnormality is evident in the regenerated antennule. These deficits are ascribed to occasional failure of regenerating neurons to reform appropriate central connections. They do not resemble the deficits following eyestalk ablation. 5. The medulla terminalis is tentatively proposed as the portion of the nervous system critical for normal antennular function that is removed by eyestalk ablation. There seem to be similarities between the effects of eyestalk ablation in the Crustacea as described here and the effects following destruction of portions of the corpora pedunculata in insects.

Oncopeltus fasciatus exhibits a facultative reproductive diapause which is triggered by short photoperiods. The duration of the triggering photoperiod (the "critical photoperiod") is a function of the environmental conditions under which populations are reared. Some individuals are apparently sensitive to critical photoperiods in the early instars, but maximum sensitivity, indicated by 100% diapause, occurs in the late 5th instar during development of the pharate adult. Sixty per cent retain sensitivity into the first five days of adulthood. Temperatures of 27°C can prevent diapause. Once initiated, diapause can be broken either by raising the temperature or increasing the day length; it is broken abruptly, for once oviposition starts, it proceeds at maximum rate. Diapause contributes significantly to a migratory strategy in the temperate parts of the range of O. fasciatus and probably does so in the tropics as well.

Physiological and ecological results from a variety of species are consistent with what seem to be valid general statements concerning insect migration. These are as follows: (i)During migration locomotory functions are enhanced and vegetative functions such as feeding and reproduction are suppressed. (ii) Migration usually occurs prereproductively in the life of the adult insect (the oogenesis-flight syndrome). (iii)Since migrant individuals are usually prereproductive, their reproductive values, and hence colonizing abilities, are at or near maximum. (iv) Migrants usually reside in temporary habitats. (v)Migrants have a high potential for population increase, r, which is also advantageous for colonizers. (vi)Both the physiological and ecological parameters of migration are modifiable by environmental factors (that is, phenotypically modifiable)to suit the prevailing conditions. Taken together, these criteria establish a comprehensive theory and adumbrate the basic strategy for migrant insects. This basic strategy is modified to suit the ecological requirements of individual species. Comparative studies of these modifications are of considerable theoretical and practical interest, the more so since most economically important insects are migrants. No satisfactory general statements can as yet be made with respect to the genotype and migration. Certainly we expect colonizing populiations to possess genotypes favoring a high r, but genotypic variation in r depends on the heritabilities of life table statistics, and such measurements are yet to be made (10, 53). The fact that flight duration can be increased by appropriate selection in Oncopeltus fasciatus, and the demonstration of additive genetic variance for this trait in Lygaeus kalmii, suggest that heritability studies of migratory behavior would also be worth pursuing. Most interesting of course, will be possible genetic correlations between migration and life history parameters. Also, migration often transports genotypes across long distances with considerable mixing of populations. An understanding of its operation therefore carries with it implications for population genetics, zoogeography, and evolutionary theory. Finally, at least parts of the above general theory would seem to be applicable to forms other than insects. Bird and insect migrations, for example, are in many respects ecologically and physiologically similar. Birds, like insects, emphasize locomotory. as opposed to vegetative functions during long-distance flight; the well-known Zugenruhe or migratory restlessness is a case in point. Further, many birds migrateat nigt at a time when they would ordinarily roost(vegetative activity). Because their life spans exceed single seasons, bird migrants are not prereproductive in the same sense that insect migrants are, and hence reproductive values do not have the same meaning(but note that some insects are also interreproductive migrants). The situaion is complicated further by the fact that in many birds adult survivorship is virtually independent of age so that colonizing ability tends to be also (10, 54). Nevertheless, birds arrive on their nesting grounds in reproductive condition with the result that migration is a colonizing episode. It is also phenotypically modifiable by environmental factors, some of which, for example, photoperiod, influence insects as well (55). The similarities between birds and insects thus seem sufficient to indicate, at least provisionally, that the theory developed for insects applies also to birds with appropriate modifications for longer life span and more complex social behavior; comparisons between insects and fish (56) lead to the same conclusion. In birds especially, and also in other forms, various functions accessory to migration such as reproductive endocrinology, energy budgets, and orientation mechanisms have been studied extensively (55, 56). But there is need in vertebrates for more data andtheoy on the ecology and physiology of migratory behavior per se in order tobetter understand its evolution and its role in ecosystem function (5, 57). Migration in any animal cannot be understood until viewed in its entirety as a physiological, behavioral, and ecological syndrome.

Electrical activity in response to light stimuli was recorded from the brain of the cricket (Gryllus domesticus) using stainless steel microelectrodes. Four basic types of elements were observed as follows: (1) units which registered ambient light intensity by frequency of firing as well as responding with transient changes in rate to stepwise increases or decreases in intensity; (2) units which fired at a higher frequency in dark than in light; (3) units which fired continuously at low level in light and responded with a transient high frequency burst to light off; and (4) units which responded with a brief burst to on and off, but tended to be “on-dominant” or “off-dominant.” Also observed were synchronized spikes in mushroom body responding primarily to light off, but also on occasion to light on, and often accompanied by single unit responses. The units registering intensity are probably homologous with units showing similar properties recorded from the visual systems of several other arthropods and usually referred to as “sustaining units.” On-off, off, and dark units are also known from other forms. The mushroom body light responses were similar to synchronized spikes recorded in cockroach mushroom body following antennal stimulation.

(1) Guild structure of the arthropod community associated with early succession from bare ground at an Old World site, southern England, and a New World site, U.S.A., was analysed from monthly samples of abundance over a 2-year period. (2) Six guilds were recognized, employing the criteria of Moran & Southwood (1982). (3) Both sites were rapidly colonized by all guilds, but the phytophages and 'tourists' were dominant. (4) Generally, the abundance (number m-2) of arthropods in all guilds was greater at the Old World site.