Spatial and temporal patterns of dispersal of western flower thrips (Thysanoptera: Thripidae) in nectarine orchards in British Columbia

Thrips were sampled from six nectarine orchards in the Dry Central Interior, British Columbia, Canada, between April and June 1993 using yellow sticky cards on posts spaced around the perimeter of each orchard. Although 12 identified species of thrips were captured, >90% of individuals were the western flower thrips, Frankliniella occidentalis (Pergande). The flight patterns and abundances of western flower thrips were compared between orchards located in two differently oriented valleys (N-S and E-W) and between orchards located close to or far from areas of wild land. Results indicate that densities of western flower thrips entering orchards, and their direction of movement, were related more to the external vegetation than either location within the two different valleys or general wind flow patterns. Western flower thrips tended to move into orchards close to ground level in early spring (late April and early May) but flew higher as ground cover grew taller and temperatures increased. Densities of western flower thrips at ground level were highest in an orchard with the densest dandelion ground cover. We conclude that the location of nectarine orchards in relation to wild areas is a major determinate of western flower thrips densities.     

Evaluation of sampling methodology for determining the phenology, relative density, and dispersion of western flower thrips (Thysanoptera: Thripidae) in nectarine orchards

Western flower thrips, Frankliniella occidentalis (Pergande), cause serious economic damage to nectarines in the Okanagan and Similkameen Valleys, British Columbia, Canada. We evaluated several sampling methods for western flower thrips for their precision and ability to predict general population trends. Beating of branches, flicking of buds, and visual estimation methods were not accurate for estimating numbers of thrips in nectarine buds. Thrips caught on sticky cards indicated general population trends, but were less efficient than collecting nectarine buds and counting thrips. Searching for thrips from buds in the field underestimated the density of both adults and larvae, and for adults, underestimated the proportion of the pale morph of western flower thrips. Dispersion patterns of thrips populations among orchards were either random or aggregated dependent on the development stage of the nectarine buds.     

A new Y-tube olfactometer for mosquitoes to measure the attractiveness of host odours

The behavioural responses of flying western flower thrips (Frankliniella occidentalis Pergande) (Thysanoptera: Thripidae) to the colour yellow and the odour anisaldehyde were examined. In a wind tunnel, upwind flight by female thrips was common in an airflow of 0.11 m s^–1 but was impeded at 0.22 m s^–1. In the absence of anisaldehyde, flying female thrips exhibited an oriented response towards a yellow cue in the wind tunnel at a wind speed of 0.11 m s^–1. The main response of females to anisaldehyde in the wind tunnel was flight inhibition. There was no evidence of an odour-induced visual response, an odour-induced anemotactic response or chemotaxis by female thrips to anisaldehyde in wind tunnel bioassays, but chemokinesis was implicated. With a matrix of yellow or black water traps with and without anisaldehyde in a greenhouse sweet pepper crop, yellow traps with anisaldehyde caught more thrips adults than yellow traps without anisaldehyde, black traps with anisaldehyde and black traps without anisaldehyde (1.3, 28 and 721 times for males respectively and 2.4, 9 and 117 times for females, respectively). Differences between respective traps were statistically significant in almost all cases. Trapping experiments using a centre-baited trap design to reduce the interaction of anisaldehyde between baited and unbaited traps were undertaken in tomato and sweet pepper greenhouse crops. When the spatial distribution of the thrips adult population within the greenhouse was taken into account, yellow water traps with anisaldehyde caught between 11 and 15 times more female and 3 and 20 times more male F. occidentalis adults than yellow traps without anisaldehyde.     

Gefiedervariationen beim M?usebussard (Buteo buteo) in Nordbayern

Zusammenfassung Die durch Linientaxierung von 1967/78 ermittelte relative Häufigkeit/Jahr von Gefiedervariationen (=5 Morphen) beim Mäusebussard in Nordbayern ergab: 0,02 % Morphe 1; ca. 98 % Morphe 2; 0,5 % Morphe 3; 0,8 % Morphe 4 und ca. 1 % Morphe 5. Helle Morphen (3–5) brüteten nicht im Beobachtungsgebiet; sie zeigten ausgeprägtes Zugverhalten bzw. Winterflucht. Die Beziehungen der Morphen 2–5 zu Temperaturen/Monat, zu strengen/milden Wintern (Mitteleuropa) und in ihrer jahreszeitlichen Verteilung sind in Tab. 2 dargestellt.

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Plumage variations in the Buzzard (Buteo buteo) in Northern Bavaria

Summary In Northern Bavaria (West-Germany) five morphs of the Buzzard could be found. Morph 1: under- and upperparts, incl. head, wings and tail, dark, without clearly visible pattern; morph 2: upperparts more or less uniform dark, pattern on underparts; morph 3: upperparts like 2, pattern on pale underparts strongly reduced or lacking; morph 4: upperparts with very large pale parts, underparts like 3; morph 5: upper- and underparts, without pattern, extremely light (primary tips are always dark, distal bar [s] on tail more or less pigmented). In 1967/78 the relative abundance/year in the differend morphs varied considerably: 0.02 % morph 1, c. 98 % morph 2, 0.5 % morph 3, 0.8 % morph 4 and c. 1 % morph 5 (roadside census). Only morph 2 was found in the breeding population. In pale morphs weather movements were common. The abundance of the morphs 2–5 showed (highly) significant correlations with winter temperatures and through phenology.

     

Brown tawny owls moult more flight feathers than grey ones

The mechanisms by which melanin‐based colour polymorphism can evolve and be maintained in wild populations are poorly known. Theory predicts that colour morphs have differential sensitivity to environmental conditions. Recently it has been proposed that colour polymorphism covaries genetically with intrinsic and behavioural properties. Plumage moult is a costly and crucial somatic maintenance function in birds. We used a long‐term data set consisting of 761 observations on 307 individuals captured between 1985 and 2010 to examine differences in partial flight feather moult between grey (pale) and brown (pheomelanic dark) colour morphs of the tawny owl. We find that the brown morph consistently moult more primary flight feathers than the grey morph whereas there is no clear difference between colour morphs in the moulting of secondary feathers. Contrary to expectations, the difference in the number of moulted flight feathers between the morphs was independent of environmental conditions, as quantified by the abundance of prey. We discuss the potential physiological and behavioural causes for and costs of the observed difference in maintenance functions between colour morphs.     

Population dynamics of western flower thrips (Thysanoptera: Thripidae) in nectarine orchards in British Columbia

Development of a control strategy for thrips attacking nectarine trees depends on an understanding of their phenology, distribution, and life history as related to characteristics of nectarine orchards. To this end, we compared the overwintering behavior, distribution, and abundance of western flower thrips, Frankliniella occidentalis (Pergande), among 11 nectarine orchards located in the dry central interior of British Columbia, Canada, during 1993 and 1994. Western flower thrips emerged from areas not previously used for agriculture (wild areas) and from within orchards before trees were out of dormancy. Flight of thrips within and around orchards peaked during early bud development, with a second major peak several weeks later after husk fall as the next generation emerged. Orchards protected from wild areas by other orchards had the lowest densities of thrips in buds. Density estimates of western flower thrips on trees were not affected by location of trees within orchards or buds within trees, but most thrips were found in the most developed buds on a tree at any one time. Thrips were not found within buds until petal was first visible on the buds. Larval feeding on buds at early petal fall resulted in serious surface russetting of fruit.     

Economic Thresholds of Western Flower Thrips (Thysanoptera: Thripidae) for Unripe Red Pepper in Greenhouse

This study was conducted to develop economic thresholds of western flower thrips (Frankliniella occidentalis Pergande) for unripe red pepper in greenhouses. To investigate the relationship between the density of thrips and resulting damages, experimental plots with five treatments (0, 4, 16, 48, 96 adults per plot) as initial release densities were established at the National Institute of Agricultural Science and Technology, Suwon, Korea, in 2004. Western flower thrips density was monitored using flower samplings and yellow sticky trap (8×13 cm) counts. Western flower thrips density was directly related to increased numbers of damaged fruits and reduced fruit yield. The number of marketable fruits produced decreased as the thrips densities increased. The major damage to pepper fruits caused by thrips was cosmetic scars that resulted from immature feeding. When flower samples or yellow trap caches were used to determine the density of thrips, which were collected on a previous sampling date, thrips densities were determined to be related to the percentage of fruits that were damaged, and a significant relationship was found for the flower samples (y = 0.3219x + 1.0792, r² = 0.8640 and for trap catches (y = 11.9209 log(x) −2.158, r² = 0.8306). The economically-tolerable ratio of damaged fruits based on control cost and market values under current greenhouse cultivation was estimated as 3.4 to 8.0%. Economic thresholds of western flower thrips ranged from 0.7 to 2.1 adults or nymphs per flower, and 2.3 to 5.7 adults per four-day sticky card count.     

On the taxonomy of Littorina africana (Mollusca: Gastropoda)

L. africana and L. knysnaensis are regarded as two morphs of a single species which exhibits a genetic cline along the south-eastern coast of southern Africa. The dark brown morph knysnaensis dominates the western, cooler end of the cline and is replaced by the pale blue morph africana at the warmer end of the cline. These conclusions are based on evidence from the latitudinal distributions, the complete range of intermediate forms regarding shell colour and shell morphology and the lack of differences in redular morphology, penial morphology or habitat.     

Origin of the disjunct distribution of flower colour polymorphism within Limonium wrightii (Plumbaginaceae) in the Ryukyu Archipelago

The sea lavender, Limonium wrightii , has six morphs of flower colour variation. The geographical distribution of flower colour morphs is disjunct; the distribution of the pink flower morph is divided into two subregions, and that of the yellow flower morph intervenes between them. The present study aimed to examine the origin of this apparent distribution pattern of flower colour in L. wrightii . Two main hypotheses (i.e. past dispersal events and phenotypic changes by natural selection and/or stochastic processes) have been proposed to account for the origin of leapfrog distribution patterns. To determine which hypothesis was applicable, we conducted a molecular phylogenetic analysis using sequence variation in chloroplast DNA (three regions of intergenic spacers, trnG - trnfM , trnV - trnM , and psbA-trnH ). We sequenced 58 accessions of L. wrightii frin 28 islands in the Ryukyu Archipelago and the Izu-Ogasawara Islands, located south of the Japanese mainland, and 12 accessions of four congeneric species. Within L. wrightii , we obtained four lineages of ten haplotypes. These lineages and haplotypes did not correlate with the different flower colours. These results indicate that the formation processes of populations are complex. The haplotypes of the pink flower morph did not show a sister relationship between the two disjunct subregions, indicating that the disjunct populations of the pink flower morphs are unlikely to share the pink flower colour as a result of common ancestry. We conclude that the observed leapfrog distribution pattern is caused by natural selection and/or stochastic processes.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 709–717.     

The pale brown allele in Cepaea nemoralis

The pale brown colour morph in Cepaea nemoralis appears to be determined by an allele at the C (colour) locus ( C ^{P B}). Pale brown is dominant to yellow, codominant with pink and recessive to dark brown. It is linked to the B locus (which controls the presence or absence of banding on the shell), but not to the U locus, which determines whether there is one band or five. In segregations of pale brown and yellow there is a significant deficiency of pale brown, suggesting that there are differences in viability between the morphs.     

Colour polymorphism and thermal capacities in Theba pisana (O.F. Müller 1774)

Several land snail species are highly polymorphic regarding their shell colouration. This polymorphism has been related to predatory effects as well as climatic reasons, assuming that dark morphs benefit from being more cryptic and therefore less prone to predation, whereas pale morphs are at an advantage under solar radiation, as they are suspected to heat up less. However, the assumption of different thermal capacities of these morphs is based on experiments with little standardisation or little environmental relevance. In this study, we aimed at measuring thermal capacities of two different morphs (pale versus dark-brown banded) of the Mediterranean land snail Theba pisana, applying a standardised and environmentally relevant test set-up, in order to prove whether darker morphs indeed do heat up more than lighter coloured morphs. We did not find any differences in the thermal capacity of the different morphs and conclude that thermal capacity of the shell is predominantly defined by its material rather than its coloration. These results are discussed with regard to previous studies on thermal characteristics of different land snail morphs and correlations between climate and morph distribution.     

The metabolic basis of life history variation: genetic and phenotypic differences in lipid reserves among life history morphs of the wing-polymorphic cricket,Gryllus firmus

The flight-capable morph of the wing-polymorphic cricket, Gryllus firmus, accumulated a substantially greater quantity of total lipid and triglyceride, compared with the obligately flightless morph, during the first five days of adulthood. Increased lipid accumulation in the flight-capable morph was genetically based, and was produced when ovarian growth is substantially reduced in that morph. Temporal changes in lipid levels suggest that the higher triglyceride reserves in the flight-capable morph fed a high-nutrient diet were produced by elevated lipid biosynthesis. By contrast, on a low-nutrient or high carbohydrate diet, increased lipid levels in the flight-capable morph appeared to result primarily from decreased lipid utilization. Increased biosynthesis or retention of triglyceride (the major flight fuel in Gryllus) by the flight-capable morph may significantly divert nutrients from egg production and hence may be an important physiological cause of its reduced ovarian growth. The obligately flightless morph allocated a greater proportion of total lipid to phospholipid than did the flight-capable morph. No functionally-significant differences in total lipid or triglyceride were produced between morphs during the last nymphal stadium. A second flightless morph, derived from the flight-capable morph by histolysis of flight muscles during adulthood, also had reduced amounts of total lipid and triglyceride but increased ovarian growth compared with the flight capable morph on the standard (high-nutrient) diet. Important qualitative and quantitative aspects of lipid metabolism differ genetically between the flight-capable and flightless morphs of G. firmus and likely contribute importantly to their respective adaptations for flight capability vs. reproduction. This is the first study to document genetically-based differences in energy reserves between morphs of a complex (phase, caste, flight) polymorphism in which morphs also differ genetically in key life history traits.     

The diurnal flight activity and influential factors of Frankliniella occidentalis in the greenhouse

Abstract The daily flight activity of western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) was examined using color, yellow-light and chemical attractant traps in a cucumber greenhouse and under controlled laboratory conditions. In the greenhouse, flying thrips were most abundant between hours 08:00–10:00, declined at mid-day, and then slightly increased during hours 14:00–16:00; however they decreased to a very low level at 18:00. The use of light traps showed no thrip flight activity during the night. The total number of thrips that flew onto cards on rainy or cloudy days was higher than that on sunny days. We reconfirmed that traps treated with attractant attracted 4.0–9.4 times more thrip than the untreated traps. Under laboratory controlled conditions, thrip flight activity was highest under light intensity between 4 000 and 6 000 lux, air temperature of 28°C, and RH of 70%. In addition, starved thrips flew more readily than non-starved thrips.     

Colour polymorphism in relation to population dynamics of the leaf beetle, Chrysomela lapponica

We studied colour morph diversity and frequencies of light and dark morphs in non-fluctuating and fluctuating populations of willow feeding leaf beetle Chrysomela lapponica in the Kola Peninsula, NW Russia. Population-specific Shannon–Weaver diversity index positively correlated with dark morph frequencies, indicating that the larger part of colour polymorphism is related with numbers and diversity of dark morphs. Among-population variation in studied characters was not explained by pollution load or predation rates, but depended on the type of the population and the stage of density change in the fluctuating populations: both colour morph diversity and frequency of dark morphs were low in declining post-outbreak populations but equally high in non-fluctuating populations and in fluctuating populations at peak densities. In time-series, both diversity index and frequency of dark morphs decreased with post-outbreak density decline in the fluctuating population, but did not change in the non-fluctuating population. In the experiment, when adults received low quality food (plants from post-outbreak site), mortality of dark morphs during the hibernation was almost doubled relative to the mortality of light morphs, whereas on high quality food the colour morphs demonstrated similar mortality. This may indicate, that decrease in colour polymorphism extent and dark morph frequencies in the declining populations is due to selective mortality of dark morphs imposed by density dependent (induced by heavy herbivore damage during an outbreak) decrease in host-plant quality (delayed inducible resistance, DIR). DIR is known as one of the factors driving herbivore populations, but our result is the first evidence that DIR may act as a factor of natural selection. Dark morphs are not only susceptible to low food quality, but also have smaller size compared to light morphs, and therefore the dark females are presumably less fecund. Thus, decrease in frequency of low-fitness (dark) individuals in post-outbreak populations and accumulation of low-fitness phenotypes at the popu-lation peak may create feedbacks contributing to regulation of density fluctuations in Ch. lapponica.     

Differential allocation of resources underlies the dispersal-reproduction trade-off in the wing-dimorphic cricket,Gryllus rubens

The cricket, Gryllus rubens (Orthoptera, Gryllidae), exists in natural populations as either a fully-winged (LW), flight-capable morph or as a short-winged (SW) morph that cannot fly. The SW morph is substantially more fecund than the LW morph. In this study we report on the physiological basis of this trade-off between flight capability and fecundity. Results from gravimetric feeding trials indicate that LW and SW morphs are equivalent in their consumption and digestion of food. However, during the adult stage, the LW morph is less efficient in converting assimilated nutrients into biomass. This may be a consequence of the respired loss of assimilated nutrients due to the maintenance of functional flight muscles in the LW morph. In both morphs the gross biomass devoted to flight muscles does not change significantly during the first 14 days of adult growth while there is a significant biomass gain in ovarian tissue mass during the same period. SW morphs have vestigial flight muscles and gain substantially more ovarian mass relative to the LW morphs. These data are consistent with a trade-off between flight muscle maintenance in the LW morph and ovarian growth in the SW form. This is the first evidence for a life-history trade-off that has a physiological basis which is limited to the allocation of acquired and assimilated nutrients within the organism.     

Coevolution of color pattern and thermoregulatory behavior in polymorphic pygmy grasshoppers Tetrix undulata

Ectothermic organisms, such as insects and reptiles, rely on external heat sources to control body temperature and possess physiological and behavioral traits that are temperature dependent. It has therefore been hypothesised that differences in body temperature resulting from phenotypic properties, such as color pattern, may translate into selection against thermally inferior phenotypes. We tested for costs and benefits of pale versus dark coloration by comparing the behaviors (i.e., basking duration and bouts) of pygmy grasshopper (Tetrix undulata) individuals exposed to experimental situations imposing a trade-off between temperature regulation and feeding. We used pairs consisting of two full-siblings of the same sex that represented different (genetically coded) color morphs but had shared identical conditions from the time of fertilization. Our results revealed significant differences in behavioral thermoregulation between dark and pale individuals in females, but not in males. Pale females spent more time feeding than dark females, regardless of whether feeding was associated with a risk of either hypothermia or overheating. In contrast, only minor differences in behavior (if any) were evident between individuals that belonged to the same color morph but had been painted black or gray to increase and decrease their heating rates. This suggests that the behavioral differences between individuals belonging to different color morphs are genetically determined, rather than simply reflecting a response to different heating rates. To test for effects of acclimation on behaviors, we used pairs of individuals that had been reared from hatchlings to adults under controlled conditions in either low or high temperature. The thermal regime experienced during rearing had little effect on behaviors during the experiments reported above, but significantly influenced the body temperatures selected in a laboratory thermal gradient. In females (but not in males) preferred body temperature also varied among individuals born to mothers belonging to different color morphs, suggesting that a genetic correlation exists between color pattern and temperature preferences. Collectively, these findings, at least in females, are consistent with the hypothesis of multiple-trait coevolution and suggest that the different color morphs represent alternative evolutionary strategies.     

Economic injury levels for western flower thrips (Thysanoptera: Thripidae) on greenhouse cucumber

Low, medium and high densities of western flower thrips, Frankliniella occidentalis (Pergande), were established in three greenhouses at the Greenhouse and Processing Crops Research Centre, Ontario, Canada, in 1996 and 1998 to develop economic injury levels for thrips on greenhouse cucumber. Thrips densities were monitored weekly using yellow sticky traps and flower counts. Fruit was harvested twice a week, graded for size, weighed, and rated for thrips damage using three damage categories. Significant yield reduction was detected 4 wk after severe fruit damage was observed in the high and medium thrips density treatments in 1996 and 7 wk in 1998. Percentage of severe damaged fruit (P(F3)) has significant linear relationships with the adult thrips density (x) that was sampled by sticky traps 1 wk before harvest (P(F3) = -0.2533 + 0.0828x) and that was sampled by flower counts 2 wk before harvest (P(F3) = -0.2025 + 0.5490x). Based on the regression equations, economic injury levels, expressed as adult thrips per sticky trap per day or adult thrips per flower, were calculated for various combinations of control costs, yield potential and fruit prices. The economic injury levels for F. occidentalis ranged from 20 to 50 adults per sticky trap per day or 3 to 7.5 per flower as determined under average greenhouse production conditions in Ontario, Canada.     

Density-dependent and frequency-dependent selection by bumblebees Bombus terrestris (L.) (Hymenoptera: Apidae)

The behaviour of bumblebee workers foraging on arrays of artificial flowers of two colour morphs was observed. Experiments were conducted on arrays of varying morph frequencies and at three different total flower densities. Bumblebees consistently showed a preference for the commonest colour morph, and this behaviour was not significantly affected by changing density. In contrast, frequency-independent preferences changed significantly with density. At low densities, there was a strong bias towards the more conspicuous colour, whereas at higher densities there was no overall colour bias. Flight distances between flowers decreased significantly at high density. Bumblebees also visited flowers of similar colours sequentially, but this behaviour was not density-dependent. It is suggested that as densities increase, there is an increased probability that bumblebees detect yellow flowers, which were probably less conspicuous compared with blue flowers, and that this might be caused by changes in flight speed with flight distance. Where there is a positive relationship between pollinator visitation and the relative fitness of a floral morph, the observed behaviour would induce positive frequency-dependent selection on a plant population with two corolla colour morphs on which the bumblebees were foraging, which would result in stabilizing selection for a single corolla colour, irrespective of density. There was no indication that rare colour morphs would be preferred at high density. The probability of different corolla colour morphs going to fixation would, however, be affected by density.     

Clinal variation in the morph ratio of Black Sparrowhawks Accipiter melanoleucus in South Africa and its correlation with environmental variables

The morph ratio distribution in polymorphic species often varies clinally, with a gradual change in morph ratios across the distributional range of the species. In polymorphic bird populations, clinal variation is rarely quantified. We describe a cline in the morph ratios of Black Sparrowhawks across South Africa, which is principally driven by a higher ratio of dark morph birds in the newly colonized southwest of the country. Across the 1400 km of our cline, the probability of a bird being a dark morph declined from over 80% close to the Cape Peninsula to under 20% in the northeast. Higher frequencies of dark morphs were associated with a higher proportion of rainfall falling during the winter breeding months. Further investigation revealed relationships between the proportion of dark morphs and altitude, amount of rainfall during the breeding months, and an interaction between this variable and temperature. These results provide some support for the suggestion that the higher frequency of dark morphs in the southwest is an adaptive response, rather than the result of a founder effect or genetic drift. These findings also suggest that, in theory, polymorphic species may be better adapted to cope with the challenges of climate change or may be able to expand their ranges more quickly into novel climatic areas, since selection pressure can act on a pre‐existing trait that may be beneficial in new conditions.