Cryptolaemus montrouzieri as a predator of the striped mealybug,Ferrisia virgata,reared on two hosts

The striped mealybug, Ferrisia virgata (Cockerell) (Hemiptera: Pseudococcidae), is a cosmopolitan pest of a variety of agricultural crops including cotton. To investigate the biological control potential of the predatory ladybird Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae) against this pest, we evaluated its developmental and reproductive fitness when feeding on F. virgata reared on pumpkin fruits or on cotton leaves and compared this to a diet of Planococcus citri Risso (Hemiptera: Pseudococcidae) reared on pumpkin fruits. F. virgata and P. citri reared on pumpkins were equally suitable prey for the pre‐imaginal stages of C. montrouzieri. Duration of total immature development was 1 day longer in C. montrouzieri offered F. virgata reared on cotton as compared with F. virgata or P. citri reared on pumpkin, whereas no significant difference was observed in survival rates. Diet significantly influenced the reproductive fitness of C. montrouzieri. Females offered P. citri reared on pumpkin had significantly shorter pre‐oviposition periods and higher fecundity and fertility than those given F. virgata reared on pumpkin or cotton leaves. F. virgata grown on cotton leaves supported the reproduction of C. montrouzieri better than F. virgata reared on pumpkin. Our study established that C. montrouzieri can successfully complete its development and reproduction when fed exclusively on F. virgata and indicates its potential as a biological control agent of this emerging cotton pest.     

Effects of three mealybug species on the development,survivorship and reproduction of the predatory lady beetle Cryptolaemus montrouzieri Mulsant

The lady beetle Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae) is an important predator of mealybugs. The development, survivorship, longevity and reproduction of C. montrouzieri feeding on three different mealybug species [Dysmicoccus neobrevipes Beardsley, Ferrisia virgata Cockerell and Planococcus minor (Maskell)] were investigated in the laboratory at 26 ± 1°C, 75-–90% RH and 14:10 (L:D) h photoperiod. Results indicated that, when feeding on different mealybugs, no significant differences were observed between developmental periods and survivorship of C. montrouzieri (from egg to adult), but differences were recorded between the sex ratios, preovipositional periods, adult longevities and reproduction of the differently treated lady beetle populations. The highest sex ratio (0.56), the longest preovipositional period (6.6 days) and adult longevity (84.8 days for females and 93.9 days for males), and the maximum fecundity (659.0 eggs/female) of C. montrouzieri were recorded when feeding on F. virgata. Moreover, C. montrouzieri had a high net reproductive rate (313.66), intrinsic rate of increase (0.0816) and finite rate of increase (1.085) when feeding on F. virgata. Results indicated that the population growth of C. montrouzieri may increase faster when feeding on F. virgata than feeding on either of the other two mealybugs.     

Effects of Transgenic Cry1Ac + CpTI Cotton on Non-Target Mealybug Pest Ferrisia virgata and Its Predator Cryptolaemus montrouzieri

Recently, several invasive mealybugs (Hemiptera: Pseudococcidae) have rapidly spread to Asia and have become a serious threat to the production of cotton including transgenic cotton. Thus far, studies have mainly focused on the effects of mealybugs on non-transgenic cotton, without fully considering their effects on transgenic cotton and trophic interactions. Therefore, investigating the potential effects of mealybugs on transgenic cotton and their key natural enemies is vitally important. A first study on the effects of transgenic cotton on a non-target mealybug, Ferrisia virgata (Cockerell) (Hemiptera: Pseudococcidae) was performed by comparing its development, survival and body weight on transgenic cotton leaves expressing Cry1Ac (Bt toxin) + CpTI (Cowpea Trypsin Inhibitor) with those on its near-isogenic non-transgenic line. Furthermore, the development, survival, body weight, fecundity, adult longevity and feeding preference of the mealybug predator Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae) was assessed when fed F. virgata maintained on transgenic cotton. In order to investigate potential transfer of Cry1Ac and CpTI proteins via the food chain, protein levels in cotton leaves, mealybugs and ladybirds were quantified. Experimental results showed that F. virgata could infest this bivalent transgenic cotton. No significant differences were observed in the physiological parameters of the predator C. montrouzieri offered F. virgata reared on transgenic cotton or its near-isogenic line. Cry1Ac and CpTI proteins were detected in transgenic cotton leaves, but no detectable levels of both proteins were present in the mealybug or its predator when reared on transgenic cotton leaves. Our bioassays indicated that transgenic cotton poses a negligible risk to the predatory coccinellid C. montrouzieri via its prey, the mealybug F. virgata.     

Morphological variation and mating compatibility within the mealybugs Pseudococcus calceolariae and P. similans (Hemiptera: Pseudococcidae), and a new synonymy

A laboratory study of isolines of Pseudococcus calceolariae (Maskell) and P. similans (Lidgett) collected from Hawke's Bay and Auckland, two widely separated regions in the North Island of New Zealand, threw doubt on the validity of the defining characters of these species. For P. similans, the number and position of oral rim tubular ducts varied widely and sometimes fell outside the defined limits for the species, and characteristic ‘stout abdominal setae’ were lost in the F₁ generation. The morphological characters that separate one species from the other were manipulated by changing the temperature at which the mealybugs developed, such that cohorts of F₁ sisters reared in the laboratory contained phenotypes of both P. calceolariae and P. similans. No impediments were found to breeding among populations of P. calceolariae and P. similans from Hawke's Bay and Auckland. All combinations of crosses between virgin females and males produced viable progeny. Those reared at 16°C laid more eggs than those reared at 23°C. The data did not suggest the existence of cryptic or sibling species, and contrasted with experiments elsewhere which quite clearly showed species incompatibility of closely related mealybugs. Examination of 160 ‘wild’ specimens of P. calceolariae from New Zealand, Australia and California (U.S.A.) and P. similans from New Zealand and Australia showed a morphological continuum from one species to the other. It is concluded that P. calceolariae and P. similans merely represent the phenotypic extremes of one widely polymorphic species, with the morphological characters of individual adults determined by the microenvironment in which they developed. Pseudococcus calceolariae is thus the senior synonym of P. similans.     

Inventory and abundance of mealybug species in immature and mature cocoa farms in Côte d'Ivoire

From 2013 to 2018, surveys were conducted in counties not previously surveyed in order to determine species of mealybugs present in the cocoa orchard in Côte d'Ivoire as well as their abundance according to the age of cocoa trees. Immature and mature cocoa trees were inspected to hand‐height in 5 and 29 counties infected with Cacao swollen shoot virus (CSSV). In each cocoa farm, mealybugs were searched for on fruits, leaves, flowers, twigs and trunks. Mealybug species were identified, and colonies were counted. Five mealybug species were identified on immature cocoa trees: Ferrisia virgata, Formicococcus njalensis, Planococcus citri, Planococcus kenyae and Pseudococcus longispinus. In addition to these species, four species, Dysmicoccus brevipes, Maconellicoccus hirsutus, Phenacoccus hargreavesi and Pseudococcus jackbeardsleyi were identified on mature cocoa trees. On immature cocoa trees, Fo. Njalensis, Pl. citri and Ps. longispinus comprised were, respectively, 35%, 33% and 19% of colonies, respectively. On mature cocoa trees, Fo. Njalensis and Pl. citri comprised 63.2% and 21.0%, and others species 15.8%. Nevertheless, the abundance of mealybug species varied according to the age of cocoa trees. The preferred organs of mealybugs were pods (74.1%) followed by twigs (13.4%) and flowers (7.4%). Previously, the mealybug Paracoccus burnerae (Brain) was found on Theobroma cacao, which is the first record for this species in Côte d'Ivoire and on this host‐plant.     

Specificity of <Emphasis Type="Italic">Anagyrus</Emphasis> sp. nov. nr. <Emphasis Type="Italic">sinope</Emphasis> and <Emphasis Type="Italic">Leptomastix dactylopii</Emphasis> for six mealybug species

In order to understand better non-target effect and potential uses, the host specificity of two parasitoid species (Anagyrus sp. nov. nr. sinope Noyes & Menezes and Leptomastix dactylopii Howard) (both Hymenoptera: Encyrtidae) for six mealybug species [Ferrisia virgata (Cockerell), Phenacoccus madeirensis Green, Phenacoccus solani Ferris, Planococcus citri (Risso),Pseudococcus longispinus (Targioni-Tozzetti) and Pseudococcus viburni (Signoret)] (all Hemiptera: Pseudococcidae) was studied through behavioral observations and laboratory rearing. The selected mealybug species represent major subfamilies and tribes of Pseudococcidae. Except for F. virgata, all mealybug species induced examinations by Anagyrus sp. nov. nr. sinope and L. dactylopii. Anagyrus sp. nov. nr. sinope was specific to P. madeirensis, which was the only mealybug species selected for oviposition and suitable for complete development of the parasitoid. No encapsulation of Anagyrus sp. nov. nr. sinope in P. madeirensis was observed. Leptomastix dactylopii accepted multiple species for oviposition, with the ranking of species preference as P. citri > P. viburni > P. longispinus > P. solani > P. madeirensis. Only P. citri, P. longispinus and P. viburni supported the development of L. dactylopii. Parasitoids developing in P. longispinus and P. viburni suffered from high encapsulation rates, while no encapsulation was observed when developing in P. citri. The results of this study suggest that Anagyrus sp. nov. nr. sinope is highly host specific. Leptomastix dactylopii, on the other hand, has a wider host range. The use of Anagyrus sp. nov. nr. sinope in a mealybug biological control program is limited to P. madeirensis and L. dactylopii to P. citri. The results presented in this study also lead us to question the accuracy of the reported host range of L. dactylopii, which include all six mealybug species tested.     

Morphological variation in Planococcoides njalensis occurring on cocoa in Ghana

A study was made to determine the extent of morphological variation in Planococcoides jalensis (Laing), the most important mealybug vector of cocoa swollen shoot virus (CSSV) in West Africa, and to resolve if more than one Planococcoides species is present. Fourteen populations of P. njalensis collected on cocoa from different localities in Ghana and one population each on Gliricidia sepium (Jacq.) Steud and Coffea canephora Pierre were compared by measured and meristic female characters using two types of morphometric analysis, Discriminant Functional Analysis and Principal Component Analysis. The analyses showed that P. njalensis exhibited a wide range of intraspecific morphological variation, particularly in meristic characters, but failed to identify new Planococcoides species. The results are discussed in relation to the control of the mealybug vectors of CSSV and to the suitability of Gliricidia as a shade tree for cocoa.     

An experimental study of morphological variation in mealybugs (Homoptera: Coccoidea: Pseudococcidae)

Abstract Cultures of a number of species of mealybug were reared under a variety of controlled environments and the effects of factors such as temperature, relative humidity and host plant on the morphology of the adult females examined. It was found that temperature had the greatest effect. The lengths of the appendages and setae, and the numbers of wax producing pores and ducts, vary over a wide range within a single species when specimens are reared at different temperatures. The relationship with temperature is generally linear, with size and pore numbers increasing with decreased temperature. However, some characters, noticeably the numbers of tubular ducts, reach their highest values at an intermediate temperature. It is suggested that while variation in size is a direct response to temperature, other characters are influenced by size or by a combination of size and environmental factors acting independently. Rearing mealybugs under different environmental conditions in order to induce morphological variation is shown to be a useful technique for resolving difficult species complexes.     

12 Phylogeny of the euglenoid loricate genera trachelomonas and strombomonas based on morphological and molecular data

Since the separation of the Trachelomonas subgroup “Saccatae” into a new genus, Strombomonas Deflandre (1930), there has been some question as to its validity. Deflandre's separation was based on morphological characteristics such as the shape of the lorica, lack of a distinctive collar, possession of a tailpiece, lack of ornamentation, and ability to aggregate particles on the lorica. Recent molecular analyses indicated that the loricate taxa were monophyletic, but few species have been sampled. The LSU rDNA from eleven Strombomonas and thirty‐eight Trachelomonas species was sequenced to evaluate the monophyly of the two genera. Bayesian and maximum‐likelihood analyses found one monophyletic clade for each genus. The Trachelomonas clade was weakly supported, but had five strongly supported subclades. Morphological characters, such as lorica development and pellicle strip reduction, also supported separation of the genera. Lorica development in Strombomonas occurred from the anterior of the cell to the posterior, forming a shroud over the protoplast whereas in Trachelomonas, a layer of mucilage was excreted over the entire protoplast, followed by creation of the collar at the anterior end. Taxa from both genera underwent exponential strip reduction at the anterior and posterior poles. In Strombomonas, only one reduction was visible in the anterior pole, while in most Trachelomonas species, two reductions were visible. Likewise, Strombomonas species possessed two whorls of strip reduction in their posterior end compared to a single whorl of strip reduction in Trachelomonas species. The combined morphological and molecular data support the retention of Trachelomonas and Strombomonas as separate genera.     

The tadpole of Scinax skuki (Anura: Hylidae) from the type locality,with a description of its larval skeleton

Herein, we provide external and internal morphological data of Scinax skuki tadpoles from its type locality. The benthic tadpole of S. skuki has eyes and nostrils positioned dorsally, vent tube dextral and reaching the free margin of the ventral fin, oral disk ventral with posterior margin concave when partially closed, labial tooth row formula 2/3, and the presence of nonpigmented spurs behind the lower jaw. These characters, together with the absence of a tectum parietale, and the shapes of the pars articularis quadrati and suprarostral, are useful for species identification and may be informative for systematic purposes.     

A new viviparous acoel Childia vivipara sp. nov. with observations on the developing embryos, sperm ultrastructure, body wall and stylet musculatures

A free‐living viviparous acoel, Childia vivipara sp. nov., from the Gullmar fjord of the Swedish coast is described. The new species is assigned to the taxon Childia based on histological, ultrastructural and molecular sequence similarities. All available molecular markers (18S rRNA, 28S rRNA and histone H3) and several morphological characters, obtained using transmission electron microscopy and confocal scanning laser microscopy of whole mount specimen stained with TRITC‐labelled phalloidin, support the placement of C. vivipara in the taxon Childia. Childia vivipara and other Childia species share the following morphological synapomorphies: well‐developed copulatory organs built of tightly packed stylet needles, proximal part of the stylet inserted into the seminal vesicle, reversed body‐wall musculature, absence of ventral diagonal muscles, presence of dorsal diagonal muscles, and presence of ventral straight longitudinal muscles between frontal pore and mouth, 9 + 1 sperm axoneme structure, six distal sperm cytoplasmic microtubules, and extensive overlap of axonemes and nucleus. The new species can be easily distinguished from other Childia species by its viviparous mode of reproduction and single curved stylet. Observations on late embryonic development based on the oldest developing embryos are discussed.     

Host specificity and comparative foraging behaviour of Aenasius vexans and Acerophagus coccois, two endo-parasitoids of the cassava mealybug

Two encyrtid parasitoids, Aenasius vexans Kerrich (Hymenoptera: Encyrtidae) and Acerophagus coccois Smith (Hymenoptera: Encyrtidae), were compared for their degree of dietary specialisation and the impact this has on their foraging strategies. Both parasitoid species are significant for biological control of the cassava mealybug, Phenacoccus herreni, Cox & Williams (Homoptera: Sternorrhyncha) a major Latin American pest of cassava, Manihot esculenta Crantz, an important root crop. Host acceptance and parasitism were analysed in seven mealybug species (with different levels of polyphagy) occurring in and around cassava fields. Results demonstrate that, in this ecosystem, An. vexans is a specialist for P. herreni while Ac. coccois is a generalist on the first and second trophic level. Of the seven mealybug species, P. herreni and P. madeirensis Green were the most acceptable hosts for Ac. coccois, followed by Ferrisia virgata Cockerell. Ac. coccois did not accept the other four mealybug species.The foraging and oviposition behaviour of individual parasitoids was observed in bioassays with cassava leaves infested by P. herreni. The two species used different strategies to locate their host. Aenasius vexans spent significantly more time walking and standing on an infested leaf and examined a host longer than did Ac. coccois. Acerophagus coccois, in contrast, spent more time for oviposition. As a consequence An. vexans parasitised more hosts in a given time than did Ac. coccois. Because the rate of offspring production of the two species did not differ, we conclude that the gregarious Ac. coccois's strategy to deposit several eggs at once might compensate for its relatively low number of ovipostitions, compared with the solitary An. vexans. These findings suggest that, given the advantages and limitations of each species, a multi-species approach to biological control of P. herreni may yield best results.     

Seed and trichome morphology of the Egyptian Fagonia (Zygophyllaceae) with emphasis on their systematic implications

Seed and trichome morphology of 13 taxa representing the genus Fagonia in Egypt have been studied using light (LM) and scanning electron microscopy (SEM). Macro‐ and micro‐morphological characters, including seed shape, seed size, seed coat sculpture, trichome density and structure were studied to evaluate their systematic significance. Seed shape and trichome density were found to have only minor taxonomic value, but seed coat sculpturing and trichome structure proved to be valuable characters. These characters offer evidence to combine certain species, e.g. Fagonia kassasii and F. schimperi in F. bruguieri, Fagonia microphylla in F. scabra and Fagonia thebaica in F. arabica. However, F. sinaica, F. tristis, F. boveana and F. isotricha should be maintained as separate species. Generally, the different patterns of seed and trichome morphology are useful in delimitation of species within the genus Fagonia, but they could not be used to characterize groups of related species.     

The past and the present through phylogenetic analysis: the rove beetle tribe Othiini now and 99 Ma

In order to classify and taxonomically describe the first two fossil Othiini (Coleoptera: Staphylinidae: Staphylininae) species from three well‐preserved specimens in Cretaceous Burmese amber, a phylogenetic analysis was conducted, combining extant and extinct taxa. A dataset of 76 morphological characters scored for 33 recent species across the subfamilies Staphylininae and Paederinae was analysed using maximum parsimony and Bayesian inference methods. The many differing phylogenetic hypotheses for higher‐level relationships in the large rove beetle subfamilies Staphylininae and Paederinae were summarized and their hitherto known fossil record was reviewed. Based on the analyses, the new extinct genus Vetatrecus gen.n. is described with two new species: V. adelfiae sp.n. and V. secretum sp.n. Both species share character states that easily distinguish them from all recent Othiini and demonstrate a missing morphological link between subfamilies Staphylininae and Paederinae. This is the first morphology‐based evidence for the paraphyly of Staphylininae with respect to Paederinae, suggested earlier by two independent molecular‐based phylogenies of recent taxa. Our newly discovered stem lineage of Othiini stresses the importance of fossils in phylogenetic analyses conducted with the aim of improving the natural classification of extant species. It also suggests that the definitions of Staphylininae and Paederinae, long‐established family‐group taxa, may have to be reconsidered. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:817F39C4-F36B-4FD9-96CD-5F8FB064C39E .     

MORPHOLOGICAL SEPARATION OF THE EUGLENOID GENERA TRACHELOMONAS AND STROMBOMONAS (EUGLENOPHYTA) BASED ON LORICA DEVELOPMENT AND POSTERIOR STRIP REDUCTION1

Since the separation of the Trachelomonas Ehrenberg subgroup “Saccatae” into a new genus, Strombomonas Deflandre (1930) , there has been some question as to its validity. Deflandre's separation was based entirely on characteristics of the lorica, including the shape of the lorica, the lack of a distinctive collar, possession of a tailpiece, lack of ornamentation, and the ability of Strombomonas species to aggregate particles on the surface of the lorica. Recent molecular analyses indicated that the loricate taxa (Trachelomonas and Strombomonas) formed a single monophyletic clade; however, the phylogenetic relationship of Strombomonas to Trachelomonas remains unclear because only two Strombomonas taxa have been sequenced to date. In this study, we evaluated the monophyly of the loricate genera using two sets of morphological characters, lorica development and pellicle strip reduction. Lorica development in Strombomonas occurred from the anterior of the cell to the posterior, forming a shroud over the protoplast. In Trachelomonas, a layer of mucilage was excreted over the entire protoplast, followed by creation of the collar at the anterior end. Taxa from both genera underwent exponential strip reduction at the anterior and posterior poles. In Strombomonas only one reduction was visible in the anterior pole, whereas in most Trachelomonas species two reductions were visible. Likewise, Strombomonas species possessed two whorls of strip reduction in the posterior end compared with a single whorl of strip reduction in Trachelomonas species. These morphological characters support the separation of Trachelomonas and Strombomonas as distinct genera.     

Phylogenetic Position of the Genus Cyrtostrombidium,with a Description of Cyrtostrombidium paralongisomum nov. spec. and a Redescription of Cyrtostrombidium longisomum Lynn & Gilron, 1993 (Protozoa,Ciliophora) Based on Live Observation,Protargol Impregnation,and 18S rDNA Sequences

We redescribe Cyrtostrombidium longisomum Lynn & Gilron, 1993, the type species of the genus Cyrtostrombidium, and describe the new species Cyrtostrombidium paralongisomum n. sp. using live observation, protargol staining and molecular data. The morphological characters of these two species are clearly distinct, i.e., dikinetid numbers in the girdle and ventral kineties; however, it is difficult to separate them by 18S rDNA sequences because they differ by only 8 bp, indicating that 18S rDNA sequences are insufficient for separating different species in the genus Cyrtostrombidium. We not only observed the position of the oral primordium in the genus Cyrtostrombidium but also observed a possibly homoplasious trait, a dorsal split in the girdle kinety, in (1) Apostrombidium, (2) Varistrombidium, and (3) Cyrtostrombidium/Williophrya. This partially supports the hypothesis of somatic ciliary pattern evolution recently put forth by Agatha and Strüder‐Kypke.     

发生在云南与东盟边境的一种新害虫——热带拂粉蚧(同翅目:粉蚧科)

【背景】热带拂粉蚧属于粉蚧类昆虫,食性杂,是危害热带和亚热带水果、蔬菜和园林植物的重要害虫,主要随水果、苗木和交通工具等介质进行远距离传播。2014年5月,在我国云南与缅甸边境发现该虫危害草本植物马松子。【方法】在收集整理热带拂粉蚧生物学、地理学等信息的基础上,介绍了其主要形态特征、寄主、分布、生物学特性,并对该有害生物的危害和防治进行了综合分析。【结果】热带拂粉蚧雌成虫长椭圆形,触角8节,体长2.5~3.0 mm,宽1.5~2.0 mm,主要通过孤雌生殖繁殖后代,形态特征与双条拂粉蚧非常相似。我国的广东、广西、海南、云南、福建等热带地区适合该虫生存和危害,一旦入侵我国会对花卉、水果和蔬菜等造成危害,给相关产业带来损失。【结论与意义】热带拂粉蚧在我国有传播扩散的可能,加强检疫是防范该虫入侵的主要手段,生物防治是治理该虫的重要措施。     

Combined phylogenetic analyses reveal interfamilial relationships and patterns of floral evolution in the eudicot order Fabales

Relationships between the four families placed in the angiosperm order Fabales (Leguminosae, Polygalaceae, Quillajaceae, Surianaceae) were hitherto poorly resolved. We combine published molecular data for the chloroplast regions matK and rbcL with 66 morphological characters surveyed for 73 ingroup and two outgroup species, and use Parsimony and Bayesian approaches to explore matrices with different missing data. All combined analyses using Parsimony recovered the topology Polygalaceae (Leguminosae (Quillajaceae + Surianaceae)). Bayesian analyses with matched morphological and molecular sampling recover the same topology, but analyses based on other data recover a different Bayesian topology: ((Polygalaceae + Leguminosae) (Quillajaceae + Surianaceae)). We explore the evolution of floral characters in the context of the more consistent topology: Polygalaceae (Leguminosae (Quillajaceae + Surianaceae)). This reveals synapomorphies for (Leguminosae (Quillajaceae + Surianaceae)) as the presence of free filaments and marginal/ventral placentation, for (Quillajaceae + Surianaceae) as pentamery and apocarpy, and for Leguminosae the presence of an abaxial median sepal and unicarpellate gynoecium. An octamerous androecium is synapomorphic for Polygalaceae. The development of papilionate flowers, and the evolutionary context in which these phenotypes appeared in Leguminosae and Polygalaceae, shows that the morphologies are convergent rather than synapomorphic within Fabales.     

CLINAL VARIATION ASSOCIATED WITH EDAPHIC ECOTONES IN HYBRID POPULATIONS OF GAILLARDIA PULCHELLA

The variety pulchella of the outcrossing annual plant species Gaillardia pulchella consists of two edaphic races in central Texas which are divergent for one morphological and four electrophoretic characters. Reduced pollen stainability in F₁ hybrids suggests the races are also divergent in chromosome structure. The recent proliferation of this species on roadsides and in pastures has led to hybridization between these races. An analysis of character variation in three hybrid populations revealed significant clinal variation associated with edaphic ecotones, and the width of these clines was found to vary among characters in a consistent pattern. It is argued that this pattern is the result of different characters experiencing different effective selection regimes, with narrower clines reflecting greater differentials in effective selection. Several mechanisms are discussed by which selection may impede the transgression of alleles across the ecotones in these populations. The results of this study are compared to the results of parallel studies on the autogamous annual species Avena barbata in California, and it is suggested that the difference between these two species in the width of clines separating edaphic ecotypes may be accounted for by their different breeding systems.