Natural flightless morphs of the ladybird beetle Adalia bipunctata improve biological control of aphids on single plants

The challenge of using ladybird beetles for biological control of insect pests such as aphids is that the adult beetles tend to fly away from the host plants. Therefore, flightless ladybirds might improve biocontrol. There are several artificial ways to obtain flightless beetles, but it may be preferable to use natural variation in flight ability. We investigated, for the first time, biocontrol by inundative augmentation of natural flightless morphs of the ladybird beetle Adalia bipunctata. Microcosm experiments using single leaves with one of three species of aphid revealed no differences in consumption behavior between flightless and winged beetles. Monitoring for 48 h of single, caged pepper plants infested with aphids of Myzus persicae nicotianae or Aulacorthum solani showed that flightless beetles had a longer residence time on the plants than winged beetles. This only translated into significantly better biocontrol of M. persicae. Despite their difference in residence time, both beetle morphs reduced the population growth of A. solani. This is probably explained by the tendency of A. solani to drop from the plant upon disturbance, and we predict that flightless beetles may outperform winged ones in the long term. Overall, our results provide a proof of principle that natural flightless A. bipunctata can improve biocontrol of aphids by ladybird beetles. However, we recognize that the effect of biocontrol will vary with the species of aphid used and that further examination in long term and large scale experiments is required.     

Morph-specific diurnal variation in allatostatin immunostaining in the corpora allata of Gryllus firmus: Implications for the regulation of a morph-specific circadian rhythm for JH biosynthetic rate

Previous studies have documented a circadian cycle in juvenile hormone (JH) biosynthesis in the long-winged, flight-capable morph, but not in the short-winged flightless morph of the cricket Gryllus firmus. One rapid and reversible inhibitor of in vitro JH biosynthesis by the corpora allata (CA) in crickets is the neuropeptide Phe-Gly-Leu/Ile-amide type of allatostatins (ASTs). To investigate the possible role of allatostatin regulation of the morph-specific circadian cycle of JH production, the quantity of this type of AST in the nerves within the CA was determined by the density of anti-AST-immunostaining in confocal images using the Image J program. The density of immunostaining was inversely related to the rate of JH biosynthesis: Immunostaining in the CA was high and did not differ between morphs early in the photophase when the in vitro rate of JH biosynthesis is low and equivalent in the morphs. However, during the end of the photophase, when the rate of JH biosynthesis rises dramatically in the flight-capable morph, but not in the flightless morph, immunostaining was significantly lower in the flight-capable compared to the flightless morph. These results indicate that morph-specific differences in delivery of AST to the CA and its probable release likely regulate the morph-specific circadian pattern of JH biosynthesis. Also, the negative correlation between AST density and JH production provides evidence for predicting the periods of altered release of these rapid-acting paracine regulators of JH biosynthesis.     

The seasonality of arboreal arthropods foraging within an Australian rainforest tree

Abstract.

• 1 The seasonality of arboreal arthropods foraging within the crown of the canopy tree Argyrodendron actinophyllum. Edlin (Sterculiaceae) was studied in a subtropical rain forest near Brisbane, Australia, during 2 years with interception traps and restricted canopy fogging.
• 2 Minimal air temperatures explained most of the seasonal variance in arthropod‘density activity'. However, the host phenology was important for several phytophagous groups. Rainfall, relative humidity and lunar phase influenced arthropod activity slightly.
• 3 Seasonal ranges were significantly different among arboreal guilds and were related to food resource availability in time, associated with both the host and the rainforest environment. Seasonal peaks of herbivores were more marked than in tropical rain forests, but less sharp than in temperate woodlands.'
• 4 Quantitative changes in arthropod density activity and abundance were important for most groups, with a marked trough during cool and dry months. The numerical contribution of most arthropod guilds to the arboreal community was not constant throughout the year.
• 5 Seasonal changes in the species composition of the arboreal community appeared not as a succession of well-defined subcommunities throughout the year, but as a continuum of species of extended seasonal ranges.
• 6 This subtropical insect-plant system presented several phenological features which were classified under tropical and temperate characteristics. This partition is open to discussion.

     

Adaptive significance of wing dimorphism in the absence of dispersal: a comparative study of wing morphs in the waterstrider, Gerris remigis

ABSTRACT.

• 1 Gerris remigis Say (Hemiptera; Gerridae) is primarily apterous, but populations with up to 33% macropters have been reported. The macropters seldom fly, and field studies have revealed no detectable differences between wing morphs in movement or survival at any time of year.
• 2 In this paper, life history traits of macropterous and apterous G. remigis are compared in an attempt to determine the mechanisms responsible for the maintenance of macroptery in this species in spite of the very low flight capacity and infrequent flight of macropters.
• 3 Development time, proportion breeding without diapause, and overwinter survival do not differ between morphs. However, pre-diapause macropterous females have a significantly shorter pre-oviposition period than apterous females. In contrast, post-diapause macropters begin reproducing later than apters, and have a lower cumulative fecundity.
• 4 These results suggest that macropters may be at a selective advantage in warm habitats which favour pre-diapause reproduction, but that apters should be favoured in the preferred, cool, lotic habitats.
• 5 However, crossing and rearing experiments indicate that wing morphology is primarily environmentally controlled in this species, and that the heritability of wing morphology is low, at best. In light of this, the relative impacts of purely phenotypic (environmental) variation, random effects, and the observed fitness differences on the maintenance of macroptery in this species are discussed.

     

Does alary dimorphism imply dispersal dimorphism in the waterstrider,Gerris remigis?

ABSTRACT.

• 1 The hypothesis that wing dimorphism reflects dimorphism for dispersal ability is tested in two populations of stream-dwelling waterstriders, Gerris remigis Say, in southern Quebec, Canada.
• 2 Movements were assessed directly by recaptures of marked adults, and indirectly by comparisons of residence times and patterns of disappearance of macropterous and apterous morphs. Sampling was done weekly, over a period of 26 months, and 4828 adult G.remigis were individually marked during this time.
• 3 Movements of >100 m were very rare for both morphs. The movement patterns of the two morphs were very similar, and differed only over the winter, when no macropters moved upstream. This latter observation suggests that macropters may be less successful than apterous individuals at moving overland, or rowing against a current at this time of year.
• 4 Residence times and patterns of disappearance of the two morphs were very similar during all seasons.
• 5 These results suggest that macropterous G.remigis do not, in general, disperse using any means unavailable to the apterous morph, and that macropters in this species should not be simply categorized as long-distance dispersers.

     

Wing and flight muscle polymorphism in a lygaeid bug, Horvathiolus gibbicollis: determinants and life history consequences

ABSTRACT.

• 1 Horvathiolus gibbicollis (Costa), a ground-living seed-feeding bug of the mediterranean region, has two wing morphs. In macrop-terous bugs both pairs of wings are fully developed. In brachypterous ones forewings are reduced to about two-thirds and hindwings to less than a third of their length in macropters.
• 2 Each morph bred true with regard to wing length when reared under variable density, food and temperature conditions for several generations.
• 3 All F₁ offspring between crosses of the two morphs were brachypterous. In F₂ approximately 25% were macropters and 75% brachypters implying monogenic control of wingform.
• 4 Flight muscles in macropters vary from fully developed to totally reduced. This variation is determined by environmental conditions during adult life. Most young adult bugs have flight muscles, and totally starved or unmated bugs retain their flight muscles. Fed and mated females histolyse flight muscles as they start laying eggs, while most males of the same group retain their muscles.
• 5 Brachypterous bugs have a smaller thorax and larger abdomen than macropterous ones.
• 6 Brachypterous bugs reach adulthood slightly before macropterous ones, and they have a distinctly shorter adult preoviposition period.
• 7 Lifetime egg production does not differ significantly between the two wing morphs. However, the temporal pattern of egg laying is different in the two morphs. The m_x-curve of macropters starts later, then attains a higher peak and finally decreases faster than that of brachypters.
• 8 Initially, macropters lay smaller eggs than brachypters, but egg volume increases with age in macropters and eventually approaches that of brachypters.
• 9 The initial increase in reproductive effort (egg volumexegg number) of macropters is concomitant with wing muscle histolysis and the mobilization of thorax space for reproduction.
• 10 Adult survival rate does not differ between the morphs.

     

Heteromorphic seeds of wheat wild relatives show germination niche differentiation

• Crop wild relatives are fundamental genetic resources for crop improvement. Wheat wild relatives often produce heteromorphic seeds that differ in morphological and physiological traits. Several Aegilops and Triticum species possess, within the same spikelet, a dimorphic seed pair, with one seed being larger than the other. A comprehensive analysis is needed to understand which traits are involved in seed dimorphism and if these aspects of variation in dimorphic pairs are functionally related.
• To this end, dispersal units of Triticum urartu and five Aegilops species were X‐rayed and the different seed morphs weighed. Germination tests were carried out on seeds, both dehulled and left in their dispersal units. Controlled ageing tests were performed to detect differences in seed longevity among seed morphs, and the antioxidant profile was assessed in terms of antioxidant compounds equipment and expression of selected antioxidant genes. We used PCA to group seed morphs sharing similar patterns of germination traits, longevity estimates and antioxidant profile.
• Different seed morphs differed significantly in terms of mass, final germination, germination timing, longevity estimates and antioxidant profile in most of the tested species. Small seeds germinated slower, had lower germination when left in their dispersal units, a higher antioxidant potential and were longer‐lived than large seeds. The antioxidant gene expression varied between morphs, with different patterns across species but not clearly reflecting the phenotypic observations.
• The results highlight different trait trade‐offs in dimorphic seeds of Aegilops and T. urartu, affecting their germination phenology and longevity, thereby resulting in recruitment niche differentiation.

     

Spatial mapping memory: methods used to determine the existence and type of cognitive maps in arboreal mammals

1. Researchers have used multiple methods to understand spatial mapping memory used by arboreal mammals for orientation: the change-point test, measures of path tortuosity, field experiments with feeding platforms, nearest-neighbour feeding tree methods, complex calculations of travel route parameters, and theoretical models.
2. This literature review provides details of all of these methods, highlights previous results from spatial mapping memory studies, and discusses perspectives for future studies.
3. Previous studies have shown that various arboreal mammals, mostly in the order Primates, can memorise spatial environments using a cognitive map. Two types of maps are characterised: the topological map, based on landmarks and reused routes, and the Euclidean map, including the ability to create shortcuts by measuring distances and distinguishing between directions. Most of the studies showed that mammals do not travel randomly but, due to the difficulty of determining which spatial map is used, the use of cognitive maps remains hypothetical.
4. When studying spatial mapping memory, data collection and analysis should account for the species’ characteristics, such as the home-range size, food preferences, and types of movements. The role of sensory cues (visual, auditory, olfactory) is crucial to understanding spatial orientation. The most relevant way to determine how arboreal mammals orientate themselves in space is by using a mix of methods: random theoretical models, collecting data in a controlled environment, measuring different parameters of travel patterns, and considering the use of sensory cues and environmental factors of the study sites.
5. Research pertaining to spatial mapping memory in arboreal mammals and forest-dwelling mammals is important for understanding cognitive abilities in mammal species, and more studies are needed in mammals of various orders.

     

The endocrine-genetic basis of life-history variation: the relationship between the ecdysteroid titer and morph-specific reproduction in the wing-polymorphic cricket Gryllus firmus

The hormonal basis of variation in life-history traits is a poorly studied topic in life-history evolution. An important step in identifying the endocrine-genetic causes of life-history variation is documenting statistical and functional associations between hormone titers and genotypes/phenotypes that vary in life-history traits. To this end, we compared the blood ecdysteroid titer and the mass of the ovaries during the first week of adulthood among a flight-capable morph and two flightless morphs of the wing-polymorphic cricket Gryllus firmus. Ecdysteroids are a group of structurally related hormones that regulate many important aspects of reproduction in insects. Both the ecdysteroid titer and ovarian mass were significantly higher in each of two flightless morphs compared with the flight-capable morph throughout the first week of adulthood. Genetically based differences in the ecdysteroid titer and ovarian mass between morphs from different selected lines were similar to phenotypically based differences among morphs from the same control (unselected) lines. By day 7 of adulthood, ovaries were typically 200-400% larger and the ecdysteroid titer was 60-300% higher in flightless versus the flight-capable morph. In addition, highly significant, positive, phenotypic correlations were observed between the ecdysteroid titer and ovarian mass in pooled samples of the two flightless and flight-capable crickets from control lines or from selected lines. The ecdysteroid titer was sufficiently elevated in the flightless morphs to account for their elevated ovarian growth. This is the first direct documentation that naturally occurring phenotypes/genotypes that differ in early fecundity, a key life-history trait, also differ phenotypically and genetically in the titer of a key reproductive hormone that potentially regulates that trait.     

Extreme adaptive life-cycle in a high arctic aphid, Acyrthosiphon svalbardicum

Abstract.

• 1 The year-round biology of a high arctic aphid is described for the first time.
• 2 The life-cycle is shown to be genetically determined, and thus markedly different to temperate species where the observed polymorphism is governed primarily by external environmental cues.
• 3 The fundatrix, which emerges from the overwintering egg, gives birth directly to sexual morphs, a phenomenon previously undescribed in the Aphidinae. This process is essentially prevented in temperate aphids by an endogenous mechanism, the interval timer.
• 4 In addition to the sexual morphs, the fundatrix produces a small number of parthenogenetic individuals (viviparae) that give rise to a third generation. This last generation consists exclusively of oviparae and males that would increase the number of overwintering eggs provided there is sufficient thermal budget for them to mature and oviposit before conditions become adverse.
• 5 The position of particular morphs in the birth sequences of the second and third generations maximize the chances of survival in harsh conditions, whilst enhancing the likelihood that individuals from the third generation will add to the number of overwintering eggs.
• 6 Guaranteed egg production combined with an in-built flexibility to produce an extra generation in particularly favourable seasons, confer adaptations to the high arctic environment, and ideally suit this aphid to exploit elevated temperatures in an era of climate change.

     

Functional dimorphic enantiostyly in monomorphic enantiostylous species of the subtribe Cassiinae (Fabaceae‐Caesalpinioideae)

• Monomorphic enantiostylous species produce flowers with a displacement of the style to the left (L) or right (R) on the same individual, and may exhibit different dynamics for the production of these floral types, which may influence levels of selfing.
• We investigated the production dynamics of L and R floral types in seven species and a variety of monomorphic enantiostylous species of the genera Senna and Chamaecrista. Our hypothesis was that most species present similar proportions of floral morphs each day. Individuals were classified daily over a period of 7 days according to the functional status, i.e. the proportion of floral morphs as functionally L, R or reciprocal (REC, i.e. similar proportions of the two floral morphs), and also according to the number of consecutive days in which they exhibited the same functional status.
• All species presented low daily flower production. Most species had individuals classified as functionally R, L and REC, and tend to repeat the same functional status over a few days, although they may change functional status during the flowering period. All species exhibited individuals that were classified as functionally reciprocal when both the daily and total number of flowers produced over 7 days was considered. The occurrence of different functional status has not yet been reported in the literature for enantiostylous species.
• The distinct strategies observed in the dynamics of floral morph production seemed likely to minimise geitonogamy and to favour cross‐pollination between individuals (xenogamy).

     

Temperature and the colour polymorphism of Philaenus spumarius (Homoptera: Aphrophoridae)

ABSTRACT.

• 1 The spittle bug, Philaenus spumarius (L.), has eleven distinct elytral colour pattern morphs in Britain, of which three are designated non-melanic and eight melanic. Frequencies of different morphs vary between populations.
• 2 The role of thermal factors in the maintenance of the balanced polymorphism was investigated by measurements of dorsal reflectance and body temperature when artifically irradiated.
• 3 Non-melanic forms had higher reflectances and showed lower temperature excesses when irradiated. There was considerable variation in reflectance for the populi–typicus morph group, and trilineatus is therefore the only true‘non-melanic’morph.
• 4 For samples along altitudinal transects in three Scottish glens, trilineatus frequencies were found to be negatively and populi-typicus and melanic frequencies positively correlated with altitude.
• 5 Thermal considerations do therefore play some role in the broad and complex selection regime of P.spumarius.

     

The African golden cat Caracal aurata: Africa's least‐known felid

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Morphological and histological examination of short‐wing formation in the winter moth Protalcis concinnata (Insecta: Lepidoptera,Geometridae)

Winter geometrid moths exhibit sexual dimorphism in wing length and female‐specific flightlessness. Female‐specific flightlessness in insects is an interesting phenomenon in terms of sexual dimorphism and reproductive biology. In the winter geometrid moth, Protalcis concinnata (Wileman), adult females have short wings and adult males have fully developed wings. Although the developmental process for wing reduction in Lepidoptera is well studied, little is known about the morphology and the developmental pattern of short‐winged flightless morphs in Lepidoptera. To clarify the precise mechanisms and developmental processes that produce short‐winged morphs, we performed morphological and histological investigations of adult and pupal wing development in the winter geometrid moth P. concinnata. Our findings showed that (a) wing development in both sexes is similar until larval‐pupal metamorphosis, (b) the shape of the sexually dimorphic wings is determined by the position of the bordering lacuna (BL), (c) the BL is positioned farther inward in females than in males, and (d) after the short pupal diapause period, the female pupal wing epithelium degenerates to approximately two‐thirds its original size due to cell death. We propose that this developmental pattern is a previously unrecognized process among flightless Lepidoptera.     

Rating the rat: global patterns and research priorities in impacts and management of rodent pests

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Juvenile hormone titer and morph-specific reproduction in the wing-polymorphic cricket,Gryllus firmus

Juvenile hormone titers and reproductive characteristics were measured in adult wing and flight-muscle morphs of the wing-polymorphic cricket, Gryllus firmus, during the first week of adulthood. This species has three morphs: one flight capable morph with fully-developed wings and fully-developed flight muscles [LW(F)], one flightless morph with fully-developed wings and histolyzed (non-functional) flight muscles [LW(H)], and another flightless morph with underdeveloped (short) wings and underdeveloped flight muscles (SW). Both flightless morphs [LW(H) and SW] had larger ovaries which contained a greater number of postvitellogenic eggs compared with the flight capable [LW(F)] morph. The juvenile hormone titer was significantly higher in SW compared with LW(F) females on days 3-7 of adulthood. On these days, the JH titer also was significantly higher in the other flightless morph, LW(H), compared with flight-capable [LW(F)] females as determined by one statistical test, but did not differ significantly by another test. The JH titer was positively correlated with ovarian mass or terminal oocyte length, but not with the number of post-vitellogenic eggs. This study is the first direct comparison of juvenile hormone titers in adult wing morphs of a wing-polymorphic insect. Results indicate that an elevated juvenile hormone titer may be at least partly responsible for one of the most distinctive features of wing-polymorphic species, the increased early fecundity of flightless females.     

Use of stable isotopes to assess the intraspecific foraging niche of males and female colour morphs of the damselfly Enallagma hageni

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Variation in singing behaviour among morphs of the sand field cricket,Gryllus firmus

1. Trade‐offs play a fundamental role in the evolution of many traits. 2. In wing‐polymorphic field crickets, the long‐winged morph can disperse from unfavourable environments, but has lower reproductive success than the short‐winged morph, because of costs associated with flight capability. 3. However, long‐winged individuals may minimise costs in favourable environments by histolysing their flight muscles and becoming flightless. 4. Few studies have examined how flight‐muscle histolysis affects male signalling and mate attraction. 5. We examined differences in singing activity and song characteristics among the flightless (short‐winged and histolysed long‐winged) and the flight‐capable male morphs, and female preferences for male song, in the sand field cricket. 6. We found: (i) both flightless morphs sang more than the flight‐capable morph, (ii) song characteristics varied among the three morphs, and (iii) females preferred songs characteristic of the long‐winged morphs. 7. Histolysis should increase mating success of long‐winged males because it increases singing activity. 8. Histolysed long‐winged males may have higher mating success than short‐winged males as they sing as frequently but produce more attractive songs. 9. Therefore, plasticity within the long‐winged morph may reduce costs of maturing in environments from which dispersal is not advantageous; non‐flying morphs may be pursuing different reproductive tactics.     

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.