Role of host-plant stimuli in sexual behaviour of small ermine moths (Yponomeuta)

ABSTRACT. 1. Studies involving experiments with olfactometers and field experiments have shown that the release of sex-pheromone by females of Yponomeuta spp. is stimulated by the presence of host plants.
2. While in Y.evonymellus L. less females emitted pheromone in the absence of host plants, in Y.padellus L. the initiation of pheromone release was delayed.
3. Females of Y.evonymellus, Y.cagnagellus Hüb., Y.padellus and Y.plumbellus D. & S. all preferred a place with host-plant odour to one with non-host odours for sex-pheromone release.
4. Male preference for host-plant odour was demonstrated in the laboratory only for Y.cagnagellus.
5. Trap catches in the field indicated a reinforcement of female attractiveness by host-plant stimuli.     

Intra- and interspecific sex-pheromone communication in the genus Yponomeuta

ABSTRACT. Male behaviour in pheromone attraction was observed in a wind tunnel for eight species of the genus Yponomeuta Latreille. Interspecific responses and the effect of some combinations of female pheromones were also investigated. The results show that the species Yponomeuta cagnagellus (Hübner), Y. malinellus Zeller, Y. rorellus (Hübner) and Y. plumbellus (Den. & Schiff.) are isolated by the specificity of their female pheromones. The other four species, Y. evonymellus (L.), Y. padellus (L.), Y. irrorellus (Hübner) and Y. vigintipunctatus (Retz.), exhibited interspecific pheromone attraction.     

Food acceptance by a monophagous and an oligophagous insect in relation to seasonal changes in host plant suitability

Food acceptance by larvae of two lepidopteran species feeding on Rosaceae, viz. Yponomeuta evonymellus (monophagous) and Y. padellus (oligophagous), was compared. The influence of seasonal changes in plants as food for both insects was examined, in particular, the effects of nitrogen and sorbitol in leaves. In the laboratory, Y. evonymellus accepts Crataegus monogyna, a host plant of Y. padellus, and Y. padellus accepts Prunus padus, the host plant of Y. evonymellus. P. padus is the most suitable food plant for Y. evonymellus. No difference in food-quality for Y. padellus was found between C. monogyna and P. padus. The performance of both species on P. padus is less influenced by seasonal changes than on Crataegus. The suitability of Crataegus decreases during the season. This is probably caused by the decrease of its nitrogen content, and not by the decrease of sorbitol in the plant. The monophagous, Y. evonymellus, is more sensitive to seasonal changes in its food when fed with a non host plant than the oligophagous Y. padellus. In oviposition experiments both species have a preference for their normal host-plants.     

Host plant use in sympatric closely related flea beetles

Studies on strategies of host plant use in sympatric-related species are significant to the theory of sympatric speciation. Altica fragariae Nakane and Altica koreana Ogloblin are sympatric closely related flea beetles found in Beijing, northern China. All their recorded host plants are in the subfamily Rosoideae of the Rosaceae, so we regard them as a model system to study interactions between herbivorous insects and plant-insect co-evolution. We conducted a set of experiments on the host preference and performance of these flea beetles to study whether these closely related species have the ability to use sympatric novel host plants and whether monophagous and oligophagous flea beetles use the same strategy in host plant use. Oviposition preference experiments showed that A. koreana, a monophagous flea beetle, displayed high host fidelity. However, A. fragariae, which is oligophagous, often made "oviposition mistakes," ovipositing on nonhost plants such as Potentilla chinensis, the host plant of A. koreana, although normal host plants were preferred over novel ones. Larval performance studies suggested that A. fragariae was able to develop successfully on P. chinensis. Feeding experiences of larvae had no effect on feeding preference, oviposition preference, and fecundity of adults. However, females were impaired in their reproductive ability when fed on nonhost plants. Therefore, A. fragariae finished their development of larval stages on P. chinensis and came back to their primary host plant, Duchesnea indica, for feeding and reproduction after eclosion.     

The role of olfactory cues in the sequential radiation of a gall‐boring beetle,Mordellistena convicta

1. Herbivorous insects often have close associations with specific host plants, and their preferences for mating and ovipositing on a specific host‐plant species can reproductively isolate populations, facilitating ecological speciation. Volatile emissions from host plants can play a major role in assisting herbivores to locate their natal host plants and thus facilitate assortative mating and host‐specific oviposition. 2. The present study investigated the role of host‐plant volatiles in host fidelity and oviposition preference of the gall‐boring, inquiline beetle, Mordellistena convicta LeConte (Coleoptera: Mordellidae), using Y‐tube olfactometers. Previous studies suggest that the gall‐boring beetle is undergoing sequential host‐associated divergence by utilising the resources that are created by the diverging populations of the gall fly, Eurosta solidaginis Fitch (Diptera: Tephritidae), which induces galls on the stems of goldenrods including Solidago altissima L. (Asteraceae) and Solidago gigantea Ait. 3. Our results show that M. convicta adults are attracted to galls on their natal host plant, avoid the alternate host galls, and do not respond to volatile emissions from their host‐plant stems. 4. These findings suggest that the gall‐boring beetles can orient to the volatile chemicals from host galls, and that beetles can use them to identify suitable sites for mating and/or oviposition. Host‐associated mating and oviposition likely play a role in the sequential radiation of the gall‐boring beetle.     

Inheritance and plasticity of adult host acceptance in Yponomeuta species: implications for host shifts in specialist herbivores

Changes in host acceptance is an important factor in the host specialization of phytophagous insects, and knowledge of the genetic organization of this behaviour is necessary in order to understand how host shifts occur. Here we describe the inheritance of adult host acceptance (oviposition) in three closely related species of Yponomeuta Latreille (Lepidoptera: Yponomeutidae), and their interspecific hybrids. Yponomeuta cagnagellus (Hübner), a specialist on Euonymus europaeus L. (Celastraceae), Y. malinellus Zeller, a specialist on Malus spp. (Rosaceae), and Y. padellus (L.), oligophagous on a number of Rosaceae, were tested for their acceptance of parental hosts in choice tests. Acceptance of E. europaeus is semi‐dominant in hybrids of Y. cagnagellus×Y. padellus, and in hybrids of Y. cagnagellus×Y. malinellus. The dominance of this acceptance was confirmed in oviposition tests with backcross hybrids: backcross hybrids F1 × Y. cagnagellus oviposited mainly on E. europaeus and F1 × Y. padellus still deposited more than half of their egg masses on E. europaeus. Reciprocal hybrids did not differ in their host acceptance, indicating that the trait is autosomal. We further studied the effect of larval food on adult host acceptance (‘Hopkins host selection principle’) in split full‐sib F1 families. Larval diet influenced oviposition only in one of two hybrid crosses. The F1 hybrid of Y. padellus× Y. cagnagellus, reared on Prunus spinosa L., deposited a significantly lower percentage of egg masses on E. europaeus compared to their full‐sib sisters fed with E. europaeus. We did not find this in the reciprocal cross. However, still more than half of the egg masses are deposited on E. europaeus by hybrids that have no experience on this host. We conclude that the semi‐dominant character of acceptance of E. europaeus and a tendency of Rosaceae‐feeding Yponomeuta to deposit egg masses on this host may have created the opportunity for the host shift of the predecessor of Y. cagnagellus from Rosaceae to the Celastraceae. This shift may have been further facilitated by a weak tendency of adults to oviposit on their larval food source.     

Development of a positive preference–performance relationship in an oligophagous beetle: adaptive learning?

The relationship between oviposition preference and larval performance is a central topic in insect–plant biology. In this study, we investigate whether the oligophagous flea beetle, Altica fragariae Nakane (Coleoptera: Chrysomelidae), exhibits a positive preference–performance relationship, and whether oviposition preference develops over time. We tested the beetles using four sympatric plant species: Duchesnea indica (Andrews) Focke (the normal host plant), Agrimonia pilosa Ledeb. (a secondary host plant), and Potentilla chinensis Ser. and Sanguisorba officinalis L. (host plants of two related Altica species) (all Rosaceae). In no‐choice experiments, both oviposition rate and offspring fitness parameters (eclosion rate, development time, and body mass) were highest on D. indica. Oviposition rate was much lower on P. chinensis than on A. pilosa, whereas offspring fitness parameters did not differ significantly between these two host plants. Offspring fitness were lowest for S. officinalis, and adult females refused to oviposit on this acceptable non‐host in a no‐choice situation. Repeated two‐choice experiments showed that the proportion of oviposition on one of the novel host plants decreased significantly over time when the alternative host plant was D. indica. In repeated two‐choice experiments using A. pilosa and P. chinensis, females mainly fed on A. pilosa but distributed their eggs equally over the two host plants, in accordance with the lack of difference in offspring fitness on those hosts. Together, these results showed that A. fragariae females develop a positive preference–performance relationship over time. We suggest that A. fragariae achieves this through adaptive learning of oviposition preference: not only does the female learn to discriminate among the host plants when there is a fitness difference for her offspring, but the female also fails to discriminate when there is no fitness difference.     

Evidence for sexual isolation as a prezygotic barrier to gene flow between morphologically divergent species of Rhagoletis fruit flies

1. Certain groups of fruit flies in the genus Rhagoletis (Diptera: Tephritidae) are exemplars for sympatric speciation via host plant shifting. Flies in these species groups are morphologically similar and overlap in their geographic ranges, yet attack different, non‐overlapping sets of host plants. Ecological adaptations related to differences in host choice and preference have been shown to be important prezygotic barriers to gene flow between these taxa, as Rhagoletis flies mate on or near the fruit of their respective host plants. Non‐host‐related assortative mating is generally absent or present at low levels between these sympatrically diverging fly populations. 2. However, some Rhagoletis taxa occasionally migrate to ‘non‐natal’ plants that are the primary hosts of other, morphologically differentiated fly species in the genus. These observations raise the question of whether sexual isolation may reduce courtship and copulation between morphologically divergent species of Rhagoletis flies, contributing to their prezygotic isolation along with host‐specific mating. 3. Using reciprocal multiple‐choice mating trials, we measured sexual isolation among nine species pairs of morphologically differentiated Rhagoletis flies. Complete sexual isolation was observed in eight of the nine comparisons, while partial sexual isolation was observed in the remaining comparison. 4. We conclude that sexual isolation can be an effective prezygotic barrier to gene flow contributing to substantial reproductive isolation between many morphologically distinct Rhagoletis species, even in the absence of differential host plant choice and host‐associated mating.     

The effects of host race, gender, and host plant distribution on alighting behavior, mating, and oviposition in Eurosta solidaginis

Eurosta solidaginis Fitch (Diptera: Tephritidae) has formed host races on Solidago altissima L. and Solidago gigantea Ait. (Asteraceae), and reproductive isolation between these host races is brought about in part by host‐associated assortative mating. Any non‐assortative mating creates the potential for gene flow between the populations, and we investigated the conditions that favored non‐assortative mating. We hypothesized that the frequency of non‐assortative mating would be influenced by differences in the behaviors of the host races and sexes and by the presence and pattern of distribution of the two host species. To test these hypotheses, we caged flies on four combinations of 32 potted host plants: all S. altissima, all S. gigantea, and cages with both host species arranged in either two pure species blocks or randomly dispersed. We recorded the number of flies of each host race that alighted on each host species and the frequency of mating within and between the host races. Males of both host races were observed on plants more frequently than females. Flies of the host race from S. gigantea (gig flies) were observed on plants in greater absolute numbers, and they mated more frequently than flies of the host race from S. altissima (alt flies). In all treatments, gig flies of both sexes were found on non‐natal host plants significantly more frequently than alt flies, and gig females showed a weaker preference for their host species than did gig males or alt flies of either gender for their respective natal hosts. Assortative mating predominated in all treatments, and flies from each host race mated more frequently in cages containing their own host plant. The frequency of non‐assortative mating varied among treatments, with the matings between alt ♀ × gig ♂ being more common in the pure S. altissima treatment and the gig ♀ × alt ♂ being more frequent in the pure S. gigantea and random treatments. Matings between gig ♂ × alt ♀ were more common overall than the reciprocal mating, because gig males were more active in pursuing matings and in alighting on the non‐natal host plant than alt flies. Non‐assortative matings were more frequent in the random than in the block treatments, but this difference was not significant. Because of strong selection against oviposition into the alternate host, we hypothesized that host plant distribution would not affect oviposition preference. We tested this hypothesis by examining the oviposition behavior of naïve, mated females in two treatments in which both host species were present: either arranged in blocks or randomly dispersed. Females oviposited only into their natal host, regardless of host plant distribution.     

Temporal variation in the genetic structure of host-associated populations of the small ermine moth Yponomeuta padellus (Lepidoptera, Yponomeutidae)

Temporal changes in allele frequencies were studied in host-associated populations of the small ermine moth Yponomeuta padellus. At one site, populations from three host plants (Sorbus aucuparia, Amelanchier larnarckii , and Crataegus spp.) were sampled annually during a four-year-period and analysed with 20 polymorphic allozyme markers. At two other sites, allele frequencies at 5- 6 enzyme loci of Y. padellus populations from two different host plants were also tested for consistency over a 13-year-pcriod. Significant allele frequency changes occurred in the short-term analysis, whereas allele frequencies remained relatively stable through time in the long-term analyses. Furthermore, allele frequencies of Y. padellus populations from Crataegus spp. were relatively stable compared to the other host populations. The role of the agents responsible for the observed patterns is discussed.     

Influence of habitat pattern on orientation during host fruit location in the tomato fruit fly, Neoceratitis cyanescens

Fruit flies have evolved mechanisms using olfactory and visual signals to find and recognize suitable host plants. The objective of the present study was to determine how habitat patterns may assist fruit flies in locating host plants and fruit. The tomato fruit fly, Neoceratitis cyanescens (Bezzi), was chosen as an example of a specialized fruit fly, attacking plants of the Solanaceae family. A series of experiments was conducted in an outdoor field cage wherein flies were released and captured on sticky orange and yellow spheres displayed in pairs within or above potted host or non-host plants. Bright orange spheres mimicking host fruit were significantly more attractive than yellow spheres only when placed within the canopy of host plants and not when either within non-host plants or above both types of plants. Additional experiments combining sets of host and non-host plants in the same cage, or spraying leaf extract of host plant (bug weed) on non-host plants showed that volatile cues emitted by the foliage of host plants may influence the visual response of flies in attracting mature females engaged in a searching behaviour for a laying site and in assisting them to find the host fruit. Moreover, the response was specific to mature females with a high oviposition drive because starved mature females, immature females and males showed no significant preference for orange spheres. Olfactory signals emitted by the host foliage could be an indicator of an appropriate habitat, leading flies to engage in searching for a visual image.     

VARIATION IN PREFERENCE AND SPECIFICITY IN MONOPHAGOUS AND OLIGOPHAGOUS SWALLOWTAIL BUTTERFLIES

Although variation in oviposition preference and specificity for host plants has been demonstrated within populations of a variety of oligophagous insect species, it is unknown whether genetic variation in host choice is lost within populations of monophagous species. Analysis of a locally monophagous butterfly species, Papilio oregonius, and a locally oligophagous species, P. zelicaon, showed significant variation in oviposition preference within populations of both species. Females of both species chose primarily their native hosts. Nonetheless, the percentages of eggs laid by individual females among the plant species and the number of plant species on which individual females laid eggs differed significantly among isofemale strains within populations. Moreover, some females within all isofemale strains of both species laid a few eggs on Foeniculum vulgare, an umbelliferous species that does not occur in the native habitats of these populations but is a host for Papilio species in other geographic areas. The results suggest that local monophagy and oligophagy in these species reflect the relative ranking among potential plant species. Both populations harbor variation in oviposition choice that could allow for host shifts if these populations invaded new habitats.     

Recognition of host-specific chemical stimulants in two sympatric host races of the pea aphid Acyrthosiphon pisum

Abstract.  1. In ecological speciation , adaptation to variation in the external environment provides the crucial push that starts the process of genetic divergence and eventually leads to speciation. This emphasis on the role of ecological specialisation in speciation events has brought with it a renewed interest in its proximate mechanisms in recently diverged groups such as host races. Here, the proximate mechanisms of feeding specialisation are investigated in two host races of the pea aphid Acyrthosiphon pisum .
2. Using alfalfa and clover extracts, enclosed in diet chambers or applied on whole plants, it is shown that feeding specialisation depends on recognition of stimulants specific to the host plant, not on deterrents or toxins specific to the non-host plants.
3. Because pea aphids mate on their host plant, feeding specialisation leads to de facto assortative mating. This study suggests that behavioural recognition of host-specific chemicals, rather than avoidance of deterrents or/and plant toxins, contributes to gene flow restriction between the alfalfa and clover host races.     

Speciation is not necessarily easier in species with sexually monomorphic mating signals

Should we have different expectations regarding the likelihood and pace of speciation by sexual selection when considering species with sexually monomorphic mating signals? Two conditions that can facilitate rapid species divergence are Felsenstein's one‐allele mechanism and a genetic architecture that includes a genetic association between signal and preference loci. In sexually monomorphic species, the former can manifest in the form of mate choice based on phenotype matching. The latter can be promoted by selection acting upon genetic loci for divergent signals and preferences expressed simultaneously in each individual, rather than acting separately on signal loci in males and preference loci in females. Both sexes in the Chrysoperla carnea group of green lacewings (Insecta, Neuroptera, Chrysopidae) produce sexually monomorphic species‐specific mating signals. We hybridized the two species C. agilis and C. carnea to test for evidence of these speciation‐facilitating conditions. Hybrid signals were more complex than the parents and we observed a dominant influence of C. carnea. We found a dominant influence of C. agilis on preferences in the form of hybrid discrimination against C. carnea. Preferences in hybrids followed patterns predicting preference loci that determine mate choice rather than a one‐allele mechanism. The genetic association between signal and preference we detected in the segregating hybrid crosses indicates that speciation in these species with sexually monomorphic mating signals can have occurred rapidly. However, we need additional evidence to determine whether such genetic associations form more readily in sexually monomorphic species compared to dimorphic species and consequently facilitate speciation.     

Oviposition,larval preference,and larval performance in two polyphagous species: does the larva know best?

It is expected that females preferentially oviposit on plant hosts that allow for optimal larval performance. However, this expectation contradicts empirical evidence where adults do not always choose the best host for their descendants. Recent evidence suggests that females’ host selection depends on the number of potential hosts. Females from oligophagous species seem to be able to choose an appropriate host in terms of larval performance, whereas in polyphagous species, adult oviposition preference is not related with larval performance. This suggests that larvae in polyphagous species could be taking a more active role in host selection than their mothers. Here, we evaluated the oviposition preference and the larval preference and performance of two polyphagous species of economic importance, Copitarsia decolora (Guenée) (Lepidoptera: Noctuidae: Cuculliinae) and Peridroma saucia (Hübner) (Lepidoptera: Noctuidae: Noctuinae), on eight species of cultivated plants. In laboratory and greenhouse choice assays, we tested adult preference for oviposition and larval preference at 1 and 24 h. Larval performance was measured in terms of survival to adulthood, length of larval period, and pupal weight. We found that both adult females and larvae actively choose their hosts and that the larval preference toward the hosts is related to the females’ preference in both herbivore species. However, the females and larvae did not preferentially select the host with the best larval performance, indicating that larval performance is not related to female or larval preference and that other selective pressures are influencing the choice of the host plant in these two species.     

桃红颈天牛成虫的交配行为及对不同寄主植物的栖落和产卵选择性

【目的】桃红颈天牛Aromia bungii是一种钻蛀性害虫,主要以蔷薇科果树和观赏树木为寄主。本研究旨在搞清楚桃红颈天牛成虫的交配产卵行为及其对不同寄主植物的栖落和产卵选择性。【方法】选择苹果Malus pumila、山樱花Prunus serrulata var. spontanea、桃Prunus persica、西府海棠Malus micromalus、毛樱桃Prunus tomentosa、秋子梨Pyrus ussuriensis、杏Prunus armeniaca和红叶李Prunus cerasifera var. atropurea 8种寄主植物木段,构建为成虫的活动场所,观察桃红颈天牛成虫的交配和产卵行为,并测定桃红颈天牛成虫在不同寄主木段上的栖落数量和产卵量。【结果】桃红颈天牛成虫的交配行为分为避让、打斗和交配3种行为。交配过程可分为相遇抱对、插入输精、交配后陪护3个阶段。桃红颈天牛雌虫产卵时无刻槽行为,喜欢将卵产于较为狭窄的缝隙中。雌虫偏好在桃树木段上栖落,雄虫则偏好在毛樱桃木段上栖落。李亚科李属的桃是桃红颈天牛最喜欢的产卵寄主木段,而雌虫在苹果亚科的苹果、海棠和秋子梨木段上产卵极少。相关性分析结果表明雌虫在寄主木段上的栖落数量与产卵量占比间存在正相关性,雄虫在寄主木段上的栖落数量与雌虫在寄主木段上的栖落数量之间也存在正相关性。【结论】桃红颈天牛雌成虫最喜欢桃树木段,雄虫对寄主植物的选择性与雌虫并不完全相同。     

Diel patterns of reproductive activities in the tomato fruit fly, Neoceratitis cyanescens

We describe the diurnal temporal and spatial patterns of reproductive activities (mating and ovipositing) of adult tomato fruit flies, Neoceratitis cyanescens (Bezzi) (Diptera: Tephritidae), under laboratory, field‐cage and field conditions. The distribution pattern of flies in the different habitats (host and non‐host plants) showed temporal, physiological and sexual differences. Mature females were observed to frequent host plants preferentially, particularly in the late afternoon. By contrast, immature females preferred to rest on non‐host plants throughout the day. Males frequented mainly non‐host plants but some were observed on host fruits in the morning. Mating activity took place preferentially on non‐host plants, in the morning. In laboratory cages, females showed a strong daily propensity to engage in mating behaviour; the mean percentage of mating females was 88 ± 5%. Time allotted to this activity (4 h 52 min ± 1 h 23 min in laboratory cages and at least 3 h for the mating pairs observed in the field) appeared considerable compared to the time required to satisfy feeding or oviposition activities. Except for a few ovipositing females observed in the morning, oviposition activities occurred mainly in the late afternoon.     

Adaptive speciation when assortative mating is based on female preference for male marker traits

Adaptive speciation occurs when frequency-dependent ecological interactions generate conditions of disruptive selection to which lineage splitting is an adaptive response. Under such selective conditions, evolution of assortative mating mechanisms enables the break-up of the ancestral lineage into diverging and reproductively isolated descendent species. Extending previous studies, I investigate models of adaptive speciation due to the evolution of indirect assortative mating that is based on three different mating traits: the degree of assortativity, a female preference trait and a male marker trait. For speciation to occur, linkage disequilibria between different mating traits, e.g. between female preference and male marker traits, as well as between mating traits and the ecological trait, must evolve. This can lead to novel speciation scenarios, e.g. when reproductive isolation is generated by a splitting in the degree of assortativeness, with one of the emerging lineages mating assortatively, and the other one disassortatively. I investigate the effects of variation in various model parameters on the likelihood of speciation, as well as robustness of speciation to introducing costs of assortative mating. Even though in the models presented speciation requires the genetic potential for strong assortment as well as rather restrictive ecological conditions, the results show that adaptive speciation due to the evolution of assortative mating when mate choice is based on separate female preference and male marker traits is a theoretically plausible evolutionary scenario.     

Divergent host plant preference causes assortative mating between sympatric host races of the ladybird beetle,Henosepilachna diekei

Divergent host preference (i.e. host fidelity) plays a significant role in the speciation process in phytophagous insects. However, how and to what extent this divergence reduces gene flow between populations has rarely been measured. Here, we estimated the intensity of assortative mating caused solely by host fidelity in two host races of the phytophagous ladybird beetle Henosepilachna diekei, specialized on Mikania micrantha (Asteraceae) and Leucas lavandulifolia (Lamiaceae) in West Java, Indonesia. These host races mated randomly in the absence of host plants under laboratory conditions, but demonstrated nearly complete assortative mating in field cages with the two host plants, by spending almost all of their time on their respective host plants. The frequency of assortative mating in the field cages was not affected drastically by host plant patch structure. These results suggest that fidelity to the different host plants yields directly almost complete reproductive isolation between the host races by limiting the habitat on the respective host plant. In addition, the high host fidelity also ensures female oviposition on the original host plant. As larvae cannot survive on non‐host plants, a positive association between female oviposition preference and larval performance on the host plant on which the beetles are specialized will further facilitate the evolution of host fidelity. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 606–614.     

Conditions for sympatric speciation: A diploid model incorporating habitat fidelity and non-habitat assortative mating

Summary Three types of genes have been proposed to promote sympatric speciation: habitat preference genes, assortative mating genes and habitat-based fitness genes. Previous computer models have analysed these genes separately or in pairs. In this paper we describe a multilocus model in which genes of all three types are considered simultaneously. Our computer simulations show that speciation occurs in complete sympatry under a broad range of conditions. The process includes an initial diversification phase during which a slight amount of divergence occurs, a quasi-equilibrium phase of stasis during which little or no detectable divergence occurs and a completion phase during which divergence is dramatic and gene flow between diverging habitat morphs is rapidly eliminated. Habitat preference genes and habitat-specific fitness genes become associated when assortative mating occurs due to habitat preference, but interbreeding between individuals adapted to different habitats occurs unless habitat preference is almost error free. However, nonhabitat assortative mating, when coupled with habitat preference can eliminate this interbreeding. Even when several loci contribute to the probability of expression of non-habitat assortative mating and the contributions of individual loci are small, gene flow between diverging portions of the population can terminate within less than 1000 generations.