Choosy mothers pick challenging plants: maternal preference and larval performance of a specialist herbivore are not linked

1. Maternal preference is a dynamic process and interactions between preference and performance are fundamental for understanding evolutionary ecology and host association in insect–plant interactions. In the present study, the hypothesis of preference–performance was tested by offering solanaceous specialist Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) larvae and adult females four plant congeners that ranged in suitability. 2. Larval feeding, development, oviposition, plant glycoalkaloids, and headspace volatiles in the four plant species were analysed to examine the extent of variation, which might explain performance–preference differences. 3. It was found that larval performance was mismatched with adult oviposition preferences. Adults laid more eggs on Solanum immite Dunal plants, which were poor hosts for larval development, feeding, and survival, compared to the other three Solanum species. 4. Chemical plant defenses, in general, did not correlate with performance or preference, but some plant volatiles may have played a role in resolving female choice. Glycoalkaloids such as solanine and chaconine were detected in similar amounts in preferred and non‐preferred hosts, but there was significantly more limonene in the headspace of S. immite than in S. tuberosum L. 5. The present findings suggest that we must consider the risk‐spreading hypothesis in cases where preference and performance are not positively correlated, particularly in specialist herbivores that can feed on a diversity of congener plants and may attempt to expand their exploits to other solanaceae species.     

Choice,performance and heritability of performance of specialist and generalist insect herbivores towards cacalol and seneciphylline,two allelochemicals of Adenostyles alpina (Asteraceae)

We compared the effects of a sesquiterpene (ST, cacalol) and a pyrrolizidine alkaloid (PA, seneciphylline), both occurring in Adenostyles alliariae, on food choice and performance of specialist and generalist insect herbivores which are all known to feed or live on A. alliariae. In choice experiments we investigated whether the compounds were preferred, deterrent or had no effect. All specialist species Aglaostigma discolor (Hymenoptera, Tenthredinidae), Oreina cacaliae (Coleoptera, Chrysomelidae) and O. speciosissima avoided feeding when confronted with the combination of compounds. Only larvae of A. discolor avoided the single ST treatment as well. Larvae of the generalist species Callimorpha dominula (Lepidoptera, Arctiidae), Cylindrotoma distinctissima (Diptera, Tipulidae) and Miramella alpina (Caelifera, Acrididae) generally avoided feeding from PA, ST and PAST treatments. The only exception were caterpillars of C. dominula which were indiscriminate towards PA when naive, and preferred to feed on the PA treatment when they had experienced the compound before. Performance, measured as the growth of larvae on the different treatments in a no choice situation over a period of 10–17 days, was not different between treatments in the specialist leaf beetles O. cacaliae and O. speciosissima. Their avoidance of the combination treatment in the choice experiments had no obvious effect on growth when forced to feed from the treatment. In the generalist C. dominula only the high concentration combination treatment (PAST) reduced growth of the larvae due to decreased consumption. In C. distinctissima we found reduced growth in all treatments except one (PA3%). Poor growth performance in C. distinctissima was due to postingestive physiological effects of all treatments and additionally to consumption reduction in high‐dose ST treatments. Genetic variability (broad sense heritability) of growth performance metabolism varied in accordance with the specialization degree of the species. O. cacaliae, the most specialized species, had no significant heritability; O. speciosissima, the less specialized specialist, had a heritability of 0.46; C. dominula, the PA adapted generalist species, had a heritability of 0.64; C. distinctissima, the generalist with no apparent adaptations, had a heritability of 0.84.     

Plant‐mediated effects of host plant density on a specialist herbivore of Solanum carolinense

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Native generalist herbivores promote invasion of a chemically defended seaweed via refuge‐mediated apparent competition

Refuge‐mediated apparent competition was recently suggested as a mechanism that enables plant invasions. The refuge characteristics of introduced plants are predicted to enhance impacts of generalist herbivores on native competitors and thereby result in an increased abundance of the invader. However, this prediction has so far not been experimentally verified. This study tested if the invasion of a chemically defended seaweed is promoted by native generalist herbivores via refuge‐mediated apparent competition. The invader was shown to offer herbivores a significantly better refuge against fish predation compared with native seaweeds. Furthermore, in an experimental community, the presence of herbivores decreased the performance of neighbouring native seaweeds, but increased growth and relative abundance of the invader. These results provides the first experimental evidence that native generalist herbivores can shift a community towards a dominance of a well‐defended invader, inferior to native species in direct competitive interactions, by means of refuge‐mediated apparent competition.     

A plant pathogen modulates the effects of secondary metabolites on the performance and immune function of an insect herbivore

Host plant chemical composition critically shapes the performance of insect herbivores feeding on them. Some insects have become specialized on plant secondary metabolites, and even use them to their own advantage such as defense against predators. However, infection by plant pathogens can seriously alter the interaction between herbivores and their host plants. We tested whether the effects of the plant secondary metabolites, iridoid glycosides (IGs), on the performance and immune response of an insect herbivore are modulated by a plant pathogen. We used the IG‐specialized Glanville fritillary butterfly Melitaea cinxia, its host plant Plantago lanceolata, and the naturally occurring plant pathogen, powdery mildew Podosphaera plantaginis, as model system. Pre‐diapause larvae were fed on P. lanceolata host plants selected to contain either high or low IGs, in the presence or absence of powdery mildew. Larval performance was measured by growth rate, survival until diapause, and by investment in immunity. We assessed immunity after a bacterial challenge in terms of phenoloxidase (PO) activity and the expression of seven pre‐selected insect immune genes (qPCR). We found that the beneficial effects of constitutive leaf IGs, that improved larval growth, were significantly reduced by mildew infection. Moreover, mildew presence downregulated one component of larval immune response (PO activity), suggesting a physiological cost of investment in immunity under suboptimal conditions. Yet, feeding on mildew‐infected leaves caused an upregulation of two immune genes, lysozyme and prophenoloxidase. Our findings indicate that a plant pathogen can significantly modulate the effects of secondary metabolites on the growth of an insect herbivore. Furthermore, we show that a plant pathogen can induce contrasting effects on insect immune function. We suspect that the activation of the immune system toward a plant pathogen infection may be maladaptive, but the actual infectivity on the larvae should be tested.     

Plant composition,not richness,drives occurrence of specialist herbivores

1. How herbivore plant diversity relationships are shaped by the interplay of biotic and abiotic environmental variables is only partly understood. For instance, plant diversity is commonly assumed to determine abundance and richness of associated specialist herbivores. However, this relationship can be altered when environmental variables such as temperature covary with plant diversity. 2. Using gall‐inducing arthropods as focal organisms, biotic and abiotic environmental variables were tested for their relevance to specialist herbivores and their relationship to host plants. In particular, the hypothesis that abundance and richness of gall‐inducing arthropods increase with plant richness was addressed. Additionally, the study asked whether communities of gall‐inducing arthropods match the communities of their host plants. 3. Neither abundance nor species richness of gall‐inducing arthropods was correlated with plant richness or any other of the tested environmental variables. Instead, the number of gall species found per plant decreased with plant richness. This indicates that processes of associational resistance may explain the specialised plant herbivore relationship in our study. 4. Community composition of gall‐inducing arthropods matched host plant communities. In specialised plant herbivore relationships, the presence of obligate host plant species is a prerequisite for the occurrence of its herbivores. 5. It is concluded that the abiotic environment may only play an indirect role in shaping specialist herbivore communities. Instead, the occurrence of specialist herbivore communities might be best explained by plant species composition. Thus, plant species identity should be considered when aiming to understand the processes that shape diversity patterns of specialist herbivores.     

Oviposition and feeding behaviour by the vine weevil Otiorhynchus sulcatus on red raspberry: effects of cultivars and plant nutritional status

• 1 The vine weevil Otiorhynchus sulcatus is a major pest of horticultural crops worldwide, with root‐feeding larvae causing most damage. Adult oviposition aboveground may therefore influence levels of damage as the larvae are relatively immobile after oviposition.
• 2 The present study investigated feeding and oviposition behaviour on red raspberry Rubus idaeus using intact plants, ensuring that choices reflected the realistic differences in cultivar appearance and chemical composition. Previous studies investigating vine weevil feeding and oviposition on other crops have used excised plant material, which may inadvertently influence behaviour.
• 3 Adult weevils significantly preferred to feed on particular cultivars in the choice experiment (e.g. Tulameen), although they consumed significantly more foliage (0.22–1.03 cm²/day) on different raspberry cultivars (e.g. Glen Moy, Glen Rosa and a wild accession) in no‐choice situations.
• 4 In choice experiments, weevils tended to avoid laying eggs on some cultivars (e.g. Glen Moy and the wild accession). The number of eggs laid (1.91–4.32 eggs per day) did not, however, differ significantly between the cultivars in a no‐choice situation. Foliar nitrogen and magnesium concentrations were positively, although weakly, correlated with the total number of eggs laid.
• 5 The present study highlights the importance of considering both choice and no‐choice tests when assessing crop susceptibility to attack because weevils may avoid feeding on certain cultivars (e.g. Glen Moy) when given a choice, although this would cause significant damage to such cultivars if they were grown in monoculture (i.e. when there is no alternative).

     

Phenotypic plasticity in chemical defence of butterflies allows usage of diverse host plants

Host plant specialization is a major force driving ecological niche partitioning and diversification in insect herbivores. The cyanogenic defences of Passiflora plants keep most herbivores at bay, but not the larvae of Heliconius butterflies, which can both sequester and biosynthesize cyanogenic compounds. Here, we demonstrate that both Heliconius cydno chioneus and H. melpomene rosina have remarkable plasticity in their chemical defences. When feeding on Passiflora species with cyanogenic compounds that they can readily sequester, both species downregulate the biosynthesis of these compounds. By contrast, when fed on Passiflora plants that do not contain cyanogenic glucosides that can be sequestered, both species increase biosynthesis. This biochemical plasticity comes at a fitness cost for the more specialist H. m. rosina, as adult size and weight for this species negatively correlate with biosynthesis levels, but not for the more generalist H. c. chioneus. By contrast, H. m rosina has increased performance when sequestration is possible on its specialized host plant. In summary, phenotypic plasticity in biochemical responses to different host plants offers these butterflies the ability to widen their range of potential hosts within the Passiflora genus, while maintaining their chemical defences.     

Variation in host plant has no effect on the performance and fitness‐related traits of the specialist herbivore Pareuchaetes insulata

The effect of host plant dissimilarity on insect preference and performance was tested using two morphological forms of Chromolaena odorata (L.) King & Robinson (Asteraceae) (one from Florida, USA, another from South Africa), and a specialist herbivore, Pareuchaetes insulata (Walker) (Lepidoptera: Erebidae: Arctiinae) from Florida, USA, that was introduced as a biological control agent in South Africa. Although this insect did establish at one site, of some 21 sites at which over 800 000 individuals were released, its population level in the field has remained low after an initial outbreak in 2006. To explain the poor performance of P. insulata, we hypothesised that P. insulata larvae prefer Floridian C. odorata to the southern African C. odorata, which is morphologically and probably genetically distinct, and that larvae reared on Floridian C. odorata should have higher fitness and performance. We tested this by comparing insect performance metrics on each of the two plant forms in laboratory experiments. Apart from pupal mass, which was significantly greater on southern African C. odorata, P. insulata performance metrics were similar on both plant forms; there were no significant differences in total leaf area consumed, egg and larval development, immature survival rates, feeding index, host suitability index, growth index, and fecundity between the Floridian and southern African C. odorata plants. In sum, we could not demonstrate that differences in plant forms in C. odorata are responsible for the poor performance of P. insulata in South Africa.     

Strong selection on mandible and nest features in a carpenter bee that nests in two sympatric host plants

Host plants are used by herbivorous insects as feeding or nesting resources. In wood‐boring insects, host plants features may impose selective forces leading to phenotypic differentiation on traits related to nest construction. Carpenter bees build their nests in dead stems or dry twigs of shrubs and trees; thus, mandibles are essential for the nesting process, and the nest is required for egg laying and offspring survival. We explored the shape and intensity of natural selection on phenotypic variation on three size measures of the bees (intertegular width, wing length, and mandible area) and two nest architecture measures (tunnel length and diameter) on bees using the native species Chusquea quila (Poaceae), and the alloctonous species Rubus ulmifolius (Rosaceae), in central Chile. Our results showed significant and positive linear selection gradients for tunnel length on both hosts, indicating that bees building long nests have more offspring. Bees with broader mandibles show greater fitness on C. quila but not on R. ulmifolius. Considering that C. quila represents a selective force on mandible area, we hypothesized a high adaptive value of this trait, resulting in higher fitness values when nesting on this host, despite its wood is denser and hence more difficult to be bored.     

Priming of antiherbivore defensive responses in plants

Defense priming is defined as increased readiness of defense induction. A growing body of literature indicates that plants (or intact parts of a plant) are primed in anticipation of impending environmental stresses, both biotic and abiotic, and upon the following stimulus, induce defenses more quickly and strongly. For instance, some plants previously exposed to herbivore‐inducible plant volatiles (HIPVs) from neighboring plants under herbivore attack show faster or stronger defense activation and enhanced insect resistance when challenged with secondary insect feeding. Research on priming of antiherbivore defense has been limited to the HIPV‐mediated mechanism until recently, but significant advances were made in the past three years, including non‐HIPV‐mediated defense priming, epigenetic modifications as the molecular mechanism of priming, and others. It is timely to consider the advances in research on defense priming in the plant–insect interactions.     

Parasitic wasp‐associated symbiont affects plant‐mediated species interactions between herbivores

Microbial mutualistic symbiosis is increasingly recognised as a hidden driving force in the ecology of plant–insect interactions. Although plant‐associated and herbivore‐associated symbionts clearly affect interactions between plants and herbivores, the effects of symbionts associated with higher trophic levels has been largely overlooked. At the third‐trophic level, parasitic wasps are a common group of insects that can inject symbiotic viruses (polydnaviruses) and venom into their herbivorous hosts to support parasitoid offspring development. Here, we show that such third‐trophic level symbionts act in combination with venom to affect plant‐mediated interactions by reducing colonisation of subsequent herbivore species. This ecological effect correlated with changes induced by polydnaviruses and venom in caterpillar salivary glands and in plant defence responses to herbivory. Because thousands of parasitoid species are associated with mutualistic symbiotic viruses in an intimate, specific relationship, our findings may represent a novel and widespread ecological phenomenon in plant–insect interactions.     

Preference for C4 shade grasses increases hatchling performance in the butterfly,Bicyclus safitza

The Miocene radiation of C₄ grasses under high‐temperature and low ambient CO₂ levels occurred alongside the transformation of a largely forested landscape into savanna. This inevitably changed the host plant regime of herbivores, and the simultaneous diversification of many consumer lineages, including Bicyclus butterflies in Africa, suggests that the radiations of grasses and grazers may be evolutionary linked. We examined mechanisms for this plant–herbivore interaction with the grass‐feeding Bicyclus safitza in South Africa. In a controlled environment, we tested oviposition preference and hatchling performance on local grasses with C₃ or C₄ photosynthetic pathways that grow either in open or shaded habitats. We predicted preference for C₃ plants due to a hypothesized lower processing cost and higher palatability to herbivores. In contrast, we found that females preferred C₄ shade grasses rather than either C₄ grasses from open habitats or C₃ grasses. The oviposition preference broadly followed hatchling performance, although hatchling survival was equally good on C₄ or C₃ shade grasses. This finding was explained by leaf toughness; shade grasses were softer than grasses from open habitats. Field monitoring revealed a preference of adults for shaded habitats, and stable isotope analysis of field‐sampled individuals confirmed their preference for C₄ grasses as host plants. Our findings suggest that plant–herbivore interactions can influence the direction of selection in a grass‐feeding butterfly. Based on this work, we postulate future research to test whether these interactions more generally contribute to radiations in herbivorous insects via expansions into new, unexploited ecological niches.     

Spatiotemporal variation in local adaptation of a specialist insect herbivore to its long‐lived host plant

Local adaptation of interacting species to one another indicates geographically variable reciprocal selection. This process of adaptation is central in the organization and maintenance of genetic variation across populations. Given that the strength of selection and responses to it often vary in time and space, the strength of local adaptation should in theory vary between generations and among populations. However, such spatiotemporal variation has rarely been explicitly demonstrated in nature and local adaptation is commonly considered to be relatively static. We report persistent local adaptation of the short‐lived herbivore Abrostola asclepiadis to its long‐lived host plant Vincetoxicum hirundinaria over three successive generations in two studied populations and considerable temporal variation in local adaptation in six populations supporting the geographic mosaic theory. The observed variation in local adaptation among populations was best explained by geographic distance and population isolation, suggesting that gene flow reduces local adaptation. Changes in herbivore population size did not conclusively explain temporal variation in local adaptation. Our results also imply that short‐term studies are likely to capture only a part of the existing variation in local adaptation.     

Plant‐mediated effects of soil nitrogen enrichment on a chemically defended specialist herbivore,Calophasia lunula

1. Nitrogen enrichment is an important driver of environmental change. In the present study, plant‐mediated effects of increased nitrogen on a specialist herbivore, Calophasia lunula Hufnagel, which sequesters antirrhinoside, an iridoid glycoside produced by its host plants, were examined. 2. Caterpillars were reared on Linaria dalmatica plants grown under low or high N treatments, and C. lunula performance traits and antirrhinoside levels were measured. Additionally, a bioassay was conducted with predatory ants to examine their response to antirrhinoside. 3. Nitrogen enrichment increased larval biomass and decreased larval antirrhinoside concentrations, but had no effect on plant iridoid glycoside concentrations or larval antirrhinoside content. Thus, differences in larval antirrhinoside concentrations were evidently a consequence of increased larval biomass. Additionally, nitrogen treatment had no effects on pupal performance or defence traits. 4. Bioassay results demonstrated a deterrent effect of antirrhinoside, but additional tests are necessary to evaluate the defensive role of this compound in insects. 5. Surprisingly, this study demonstrated little effect of a six‐fold increase in nitrogen availability on L. dalmatica iridoid glycoside concentrations or sequestration by C. lunula. Moreover, the results suggested that changes in plant primary chemistry were more important than secondary chemistry for this specialist herbivore, and that some insects may be insensitive to N enrichment.     

Determinants of growth rate on chemically heterogeneous host plants by specialist insects

Nine Passiflora species growing sympatrically in a Costa Rican rain forest were screened for the presence of alkaloids, tannins, cyanogenic compounds, saponins and cardenolides. The first three chemical classes were found to be present and the Passiflora species could be classified into five groups according to the presence/absence of each chemical class. In order to assess the effects of this chemical variability on specialist herbivores, larvae of Heliconius ismenius and H. melpomane were experimentally introduced to seven species of Passiflora host plants growing naturally in the rain forest. Survival and growth rates were subsequently monitored at 2 day intervals and were found to be independent of each other. Larval growth rate and larval survivorship were not significantly affected by cyanogenesis in the host plants, nor by the presence of alkaloids, tannins and non-tanning phenolic compounds. Larval growth rate was found to be correlated to percentage nitrogen in the host plants, but was independent of host water content and non-structural carbohydrate content. Although these insects would be classified as host-specific by most authors, the results indicate the presence of a sufficiently general detoxication apparatus to consume many different classes of potential toxins with no apparent ill effects. We conclude that Passiflora natural product diversity does not cause major deleterious effects in Heliconius growth characteristics and that these effects are not the major component maintaining the current pattern of larval feeding specialization in this plant-herbivore system.     

Experimental demography and the vital rates of generalist and specialist insect herbivores on native and novel host plants

1. Colonization success of species when confronted with novel environments is of interest in ecological, evolutionary and conservation contexts. Such events may represent the first step for ecological diversification. They also play an important role in adaptive divergence and speciation. 2. A species that is able to do well across a range of environments has a higher plasticity than one whose success is restricted to a single or few environments. The breadth of environments in which a species can succeed is ultimately determined by the full pattern of its vital rates in each environment. 3. Examples of organisms colonizing novel environments are insect herbivores expanding their diets to novel host plants. One expectation for insect herbivores is that species with specialized diets may display less plasticity when faced with novel hosts than generalist species. 4. We examine this hypothesis for two generalist and two specialist neotropical beetles (genus Cephaloleia: Chrysomelidae) currently expanding their diets from native to novel plants of the order Zingiberales. Using an experimental approach, we estimated changes in vital rates, life-history traits and lifetime fitness for each beetle species when feeding on native or novel host plants. 5. We did not find evidence supporting more plasticity for generalists than for specialists. Instead, we found similar patterns of survival and fecundity for all herbivores. Larvae survived worse on novel hosts; adults survived at least as well or better, but reproduced less on the novel host than on natives. 6. Some of the novel host plants represent challenging environments where population growth was negative. However, in four novel plant-herbivore interactions, instantaneous population growth rates were positive. 7. Positive instantaneous population growth rates during initial colonization of novel host plants suggest that both generalist and specialist Cephaloleia beetles may be pre-adapted to feed on some novel hosts. This plasticity in host use is a key factor for successful colonization of novel hosts. Future success or failure in the colonization of these novel hosts will depend on the demographic rates described in this research, natural selection and the evolutionary responses of life-history traits in novel environments.     

Interaction of specialist root and shoot herbivores of Alliaria petiolata and their impact on plant performance and reproduction

Abstract.  1. This study explored interactions of two spatially and temporally separated weevils and their impact on Alliaria petiolata (garlic mustard) survival, growth, and reproduction at different herbivore densities.
2. The root-mining weevil Ceutorhynchus scrobicollis attacks A. petiolata rosettes from October to April, and larvae complete development in May. The shoot-mining weevil Ceutorhynchus alliariae attacks bolting plants in April/May with larvae completing development in June–July. Priority effects were expected, with early attack of C. scrobicollis affecting the later attacking C. alliariae , mediated through changes in plant growth or chemistry.
3. Attack by C. scrobicollis significantly increased plant mortality and changed plant architecture, while C. alliariae only significantly reduced plant height. Attack by C. scrobicollis also increased nitrogen content of stems.
4. Root feeding by C. scrobicollis affected the feeding niche of C. alliariae , but increased stem nitrogen content did not result in increased stem miner survival. While reduced height and stem diameters as a result of C. scrobicollis attack reduced C. alliariae attack at the stem level, attack at plant level and recruitment was unaffected.
5. Weevil density had no effect on plant performance, most likely due to strong intraspecific competition, and there were no synergistic effects between the two herbivores.
6. Overall, attack by C. scrobicollis was more detrimental to A. petiolata growth, seed output, and survival than attack by C. alliariae . Consequently, C. scrobicollis has been prioritised as a potential biocontrol agent for control of A. petiolata in North America.     

Entrapped sand as a plant defence: effects on herbivore performance and preference

1. Abrasive material in the diet of herbivorous organisms comes from a variety of sources, including crystalline silica or calcium in plant tissues, accidentally ingested soil while digging or grazing, and entrapped substrate on the surfaces of plants. A wide variety of plants entrap substrate, usually with glandular trichomes. 2. A previous study demonstrated that entrapped sand provided resistance to herbivory in the field. In this study, the following questions were addressed: how does entrapped sand on Abronia latifolia (Nyctaginaceae) leaves and stems affect preference and performance of a common herbivore, the large‐bodied caterpillar Hyles lineata (Sphingidae); does this effect differ from those experienced by an internally feeding leaf miner? 3. Using a combination of experimental and observational approaches, it was found that sand comprised ～4–5% of ingested weight during normal feeding of H. lineata caterpillars. This entrapped sand caused extensive wear to their mandibles, they avoided sand‐covered plants when given the choice, and the sand negatively impacted performance metrics, including pupal weight, development time, and growth rate. In contrast, a leaf‐mining caterpillar did not have a preference for or against feeding on sandy plants. 4. These results are similar to studies on mandibular wear due to grasses, and herbivorous insects that feed on these two plant groups may have similar morphologies. It is hypothesised that increased wear potential may be a convergent solution to abrasive plants in both mammals (hypsodonty) and insects.     

Trees to treehoppers: genetic variation in host plants contributes to variation in the mating signals of a plant‐feeding insect

Community genetics research has demonstrated ‘bottom‐up’ effects of genetic variation within a plant species in shaping the larger community with which it interacts, such as compositions of arthropod faunas. We demonstrate that such cross‐trophic interactions also influence sexually selected traits. We used a member of the Enchenopa binotata species complex of treehoppers (Hemiptera: Membracidae) to ask whether male mating signals are influenced by host plant genetic variation. We reared a random sample of the treehoppers on potted replicates of a sample of host plant clone lines. We found that treehopper male signals varied according to the clone line on which they developed, showing that genetic variation in host plants affects male treehoppers' behavioural phenotypes. This is the first demonstration of cross‐trophic indirect genetic effects on a sexually selected trait. We discuss how such effects may play an important role in the maintenance of variation and within‐population phenotypic differentiation, thereby promoting evolutionary divergence.