Phylogeographic studies on natural populations of the South American fruit fly, <Emphasis Type="Italic">Anastrepha fraterculus</Emphasis> (Diptera: Tephritidae)

Anastrepha fraterculus is an important pest of commercial fruits in South America. The variability observed for morphological and behavioural traits as well as genetic markers suggests that A. fraterculus represents a complex of synmorphic species rather than a single biological species. We studied the correlation between geographical distribution and genetic variation in natural populations from Argentina and south Brazil. Fragments of the mitochondrial gene COII were sequenced in 28 individuals. The matrix of aligned sequences was phylogenetically analysed by parsimony, yielding a cladogram of haplotypes. Based on Templeton’s nested method, no clade showed any geographic pattern for the gene COII, indicating lack of significant association between haplotypic variability and geographic distribution. The analysis of nucleotide substitution distances by Neighbour-Joining algorithm showed that geographically distant populations exhibit low genetic distances. The corresponding trees clustered the populations without showing any geographic pattern. This result suggests that the populations studied are not reproductively isolated.     

To hybridize or not to hybridize: what separates two genetic lineages of the Chalk‐hill Blue Polyommatus coridon (Lycaenidae,Lepidoptera) along their secondary contact zone throughout eastern Central Europe?

As a consequence of postglacial range expansion, hybrid zones evolved where different genetic lineages met. In this study, we analysed the Chalk‐hill Blue Polyommatus coridon all along the contact zone of two expansive lineages. This zone stretches from the sandy areas of north‐eastern Germany, along the mountain ranges of the German–Czech border and throughout the eastern Alps. We studied allozymes (19 loci) of 38 populations (1542 individuals) and compared these data sets against 15 populations of the western and 15 populations of the eastern lineages and found different degrees of hybridization. Thus, the calcareous regions of Thuringia and Sachsen‐Anhalt were mostly colonized by the western lineage. The middle mountain ranges between Bavaria and Bohemia represented a strong barrier blocking further expansion and thus completely impeding hybridization in this region. More intense hybridization was detected in the populations of the eastern Alps, especially in the north‐eastern part, where the Danube most probably acted as an expansion corridor for both lineages followed by intensive hybridization. In the south‐eastern Alps, hybrid populations were mostly detected in the easternmost parts and along the larger river valley of Drava and Mur; pure western populations dominated in the other areas of this region. These results show that the degree of hybridization along a contact zone is correlated with the ecological demands of a species and the regional physical geographic circumstances. This finding was proved for the Chalk‐hill Blue in our study but is also the most likely scenario in most animal and plant species.     

Müllerian mimicry of a quantitative trait despite contrasting levels of genomic divergence and selection

Hybrid zones, where distinct populations meet and interbreed, give insight into how differences between populations are maintained despite gene flow. Studying clines in genetic loci and adaptive traits across hybrid zones is a powerful method for understanding how selection drives differentiation within a single species, but can also be used to compare parallel divergence in different species responding to a common selective pressure. Here, we study parallel divergence of wing colouration in the butterflies Heliconius erato and H. melpomene, which are distantly related Müllerian mimics which show parallel geographic variation in both discrete variation in pigmentation, and quantitative variation in structural colour. Using geographic cline analysis, we show that clines in these traits are positioned in roughly the same geographic region for both species, which is consistent with direct selection for mimicry. However, the width of the clines varies markedly between species. This difference is explained in part by variation in the strength of selection acting on colour traits within each species, but may also be influenced by differences in the dispersal rate and total strength of selection against hybrids between the species. Genotyping‐by‐sequencing also revealed weaker population structure in H. melpomene, suggesting the hybrid zones may have evolved differently in each species, which may also contribute to the patterns of phenotypic divergence in this system. Overall, we conclude that multiple factors are needed to explain patterns of clinal variation within and between these species, although mimicry has probably played a central role.     

Integrating classical and spatial multivariate analyses for assessing morphological variability in the endemic Iberian viper Vipera seoanei

Historical and ecological processes have deeply affected biogeographic patterns of animals. Studying morphological variability of species, using classical and spatial analyses, can elucidate these patterns and give insights on both processes. Morphological variability of the endemic Iberian viper Vipera seoanei is examined to identify morphological coherent groups, biogeographic patterns and the putative role of abiotic pressures in the geographic variation of morphological variation. Results from classic and spatial multivariate analyses over 27 morphometric traits for 468 specimens from the global range of the species were integrated. Classic analyses reported large morphological variability and confirmed the differentiation of two coherent groups, which are representatives of current subspecies. Spatial analyses reported a geographic gradient pattern from western Cantabrian Mountains to the rest of the study area. Areas of high morphological variability were found, and two spatial coherent groups with an integration zone were recognized. Significant spatial correlations and trends suggest that some traits could be under selection and may display adaptations to local environments. Although observed patterns can be attributed to Pleistocene climatic cycles, an adaptive diversification of the species is supported. The combination of classical and spatial multivariate analyses is a useful methodology to identify morphological patterns and infer underlying factors.     

Phenotypic plasticity and genetic isolation-by-distance in the freshwater mussel <Emphasis Type="Italic">Unio pictorum</Emphasis> (Mollusca: Unionoida)

Freshwater mussels (Unionoida) show high intraspecific morphological variability, and some shell morphological traits are believed to be associated with habitat conditions. It is not known whether and which of these ecophenotypic differences reflect underlying genetic differentiation or are the result of phenotypic plasticity. Using 103 amplified fragment length polymorphism (AFLP) markers, we studied population genetics of three paired Unio pictorum populations sampled from two different habitat types (marina and river) along the River Thames. We found genetic differences along the Thames which were consistent with a pattern of isolation by distance and probably reflect limited dispersal via host fish species upon which unionoid larvae are obligate parasites. No consistent genetic differences were found between the two different habitat types suggesting that morphological differences in the degree of shell elongation and the shape of dorso-posterior margin are caused by phenotypic plasticity. Our study provides the first good evidence for phenotypic plasticity of shell shape in a European unionoid and illustrates the need to include genetic data in order properly to interpret geographic patterns of morphological variation.     

Leaf morphological responses to variation in water availability for plants in the <Emphasis Type="Italic">Piriqueta caroliniana</Emphasis> complex

Distribution of plants and the expression of traits associated with environmental variation can be affected by both average conditions and the variance in conditions including extreme climatic events. We expect that these same factors should affect the distribution of plants in hybrid zones between ecologically distinct species where the hybrids should occupy ecotones or intermediate habitats. We evaluated water availability and leaf morphological differences among parental and hybrid populations of herbaceous perennial plants in the Piriqueta caroliniana complex along environmental gradients in Southeastern North America. We focus on two taxa in this group; the viridis morphotype, which occurs in southern Florida, and the caroliniana morphotype, which is distributed from northern Florida to southern Georgia. Advanced-generation hybrid derivatives of these morphotypes occupy a broad geographic region that extends across much of central Florida. Overall, we found that hybrid populations occurred in significantly drier locations, indicating that their habitat requirements are transgressive (i.e., exceeding parental values) rather than intermediate to the parental morphotypes. Water availability differed between the two sampling years, and plants displayed morphological changes in response to these changes in moisture. During the drier year, leaves were narrower and more hirsute, corroborating experimental results that these leaf traits are plastic, and confirming that plasticity occurs in natural habitats. Hybrids exhibited intermediate leaf traits (shape and size) across both years, and displayed transgressive (hair density) leaf traits during the drier year. The apparent canalization of the hybrids’ leaf morphological traits may contribute to their tolerance of variable environmental conditions and may partially explain why they have displaced the caroliniana morphotype in central Florida.     

Geographic structure of genetic and phenotypic variation in the hybrid zone between Quercus affinis and Q. laurina in Mexico

Analyzing the structure of hybrid zones is important for inferring their origin, dynamics and evolutionary significance. We examined the geographic structure of phenotypic and genetic variation in the contact zone between two Mexican red oaks, Quercus affinis and Q. laurina. A total of 105 individuals from seven populations were sampled along a 600‐km latitudinal gradient representing the distribution area of the two species and their contact zone. Individuals were genotyped for nine nuclear and four chloroplast DNA microsatellite loci (ncSSR and cpSSR, respectively), and characterized for several leaf and acorn traits. The cpSSR data revealed extensive haplotype sharing among populations of the two species, while a Bayesian assignment analysis based on ncSSRs identified two main genetic groups, each corresponding to one of the species, and two populations in the contact zone showing evidence of admixture. The proportion of genetic ancestry in the populations was strongly associated with latitude and showed a pattern of variation with the shape of a narrow sigmoidal cline. The variation in three of the seven phenotypic traits was partially congruent with molecular variation, while the other traits did not conform to a geographic cline but instead were correlated with environmental variables. In conclusion, the hybrid zone between the two oak species has some of the characteristics of a tension zone, but heterogeneous variation across traits suggests differential introgression and the action of extrinsic selection.     

Differentiation in morphology and habitat partitioning of genetically characterized Gammarus fossarum forms (Amphipoda) across a contact zone

Evolutionary processes do not affect all character types equally. Depending on environmental effects, morphological and ecological evolution may be uncoupled from molecular evolution. A natural contact zone between two forms of the amphipod Gammarus fossarum was used to assess the levels of differentiation in genetic, morphological and habitat characters. About 50 individuals per population were analysed for six enzyme loci in 72 populations and four size-standardized morphometric traits in a subset of 32 populations. Nine environmental parameters per population were used to characterize the habitat. All three character sets revealed significant divergences between G. fossarum forms. The eastern form A of G. fossarum differs from the western form B by longer setae at the second antennae (males) and longer exopodits at die third uropods (males and females). The variance partitioning within populations, among populations and between forms, was compared within the three character sets. The relative variance component between forms in genetic characters appeared to be 10-fold greater in comparison to morphological traits, indicating morphological stasis during speciation. The inter-form environmental differentiation was shown to be low, but consistent between pure populations. The eastern form A seems to prefer higher altitudes, a substrate type of gravel, stones and leaves, shading, and water with higher conductivity and pH man the western form B.     

Morphological variability of Consolida regalis seeds of south-eastern and central Europe

The aim of the paper was to analyse the morphological variability of Consolida regalis Gray (common name Forking Larkspur) seeds in populations derived from different geographic origins. A review of available literature showed scarcity of data on seed sizes and a complete lack of information describing the morphological variation of C. regalis seeds in the geographic gradient. Therefore, the main aim of the study was to analyse the morphological variability of C. regalis seeds from south-eastern and central Europe. Larkspur seeds, collected during expeditions spanning 2012–2014 from 29 sites located in Bulgaria, Romania, Moldova and Poland, were the study material. Four biometric traits were analysed, that is, the length, the width, the perimeter and the area of the seeds. The pictures of the surface structure and sculpture of the seeds were made using the scanning electron microscope (SEM). The results showed differences in biometric traits and the seed sculpture across the latitudinal transect. There was a trend observed of an increase in the length, the width, the perimeter and the area of the seeds towards the northern border of the geographic species range. Besides the geographic factor, the morphological variation of larkspur seeds was affected by environmental conditions (mainly temperature) in which plants grew and availability of habitat resources.     

Morphometric and meristic diversity of the species Scolopendra cingulata Latreille, 1829 (Chilopoda: Scolopendridae) in the Mediterranean region

Scolopendra cingulata Latreille, 1829 is the commonest scolopendromorph species in the Mediterranean area. However, its morphological and geographic variation has not been examined so far, and therefore robust hypotheses about the factors that have shaped that variability are lacking. We examined, using multivariate methods for 19 morphometric and meristic characters, the morphological variation of that species using a comprehensive sample of 503 adult specimens from 130 restricted geographic localities. The localities were distributed in three major geographic areas (Balkans, Asia Minor-Middle East, and Italy) in order to discern biological entities and to estimate the morphological relationships between populations and geographic regions. Results showed significant differences between the three geographic groups. Characters such as the distance between the paramedian sulci of the 7th and 8th tergite, the number of antennal segments, the number of spines on the dorsal side of 21st prefemur and number of spines on the 21st prefemoral process significantly discriminated populations of S. cingulata along a west-east geographic gradient. Both eastern (Balkan) and easternmost populations (Asia Minor-Middle East) showed higher mean values of antennal articles and spines of 21st prefemur than the Italian populations. No significant morphological variation was discovered between the sexes of S. cingulata apart from cephalic width and distance between the two paramedian sulci of the 7th and 8th tergite. Based on certain morphometric differences among the three major geographic regions, we suggest that S. cingulata originated in the east and colonized south European area from two different geographic routes. It is suggested that west-east morphological trend of S. cingulata is related either to the prominent palaeogeographic events of the area or to the competition with S. oraniensis Lucas, 1846 in the western Mediterranean.     

A genetically distinct hybrid zone occurs for two globally invasive mosquito fish species with striking phenotypic resemblance

Hybrid zones allow for the investigation of incipient speciation and related evolutionary processes of selection, gene flow, and migration. Interspecific dynamics, like competition, can impact the size, shape, and directional movement of species in hybrid zones. Hybrid zones contribute to a paradox for the biological species concept because interbreeding between species occurs while parental forms remain distinct. A long‐standing zone of intergradation or introgression exists for eastern and western mosquito fish (Gambusia holbrooki and G. affinis) around Mobile Bay, AL. The region has been studied episodically, over decades, making it perfect for addressing temporal dynamics and for providing a deeper understanding of the genetics of these periodically reclassified fishes (as species or subspecies). We used six microsatellite markers to assess the current population structure and gene flow patterns across 19 populations of mosquito fish and then compared our results with historical data. Genetic evidence demonstrates that the current hybrid zone is located in a similar geographic region as the historical one, even after three decades. Hybrid fish, however, demonstrate relatively low heterozygosity and are genetically distinct from western and eastern mosquito fish populations. Fin ray counts, sometimes used to distinguish the two species from one another, demonstrate more eastern (G. holbrooki) phenotype fish within the molecular genetic hybrid zone today. Mosquito fish are globally invasive, often found on the leading edge of flooded waters that they colonize, so the impact of hurricanes in the wake of climate change was also evaluated. An increase in the frequency and intensity of hurricanes in the hybrid region has occurred, and this point warrants further attention since hurricanes are known to move these aggressive, invasive species into novel territory. This work contributes to our classical understanding of hybrid zone temporal dynamics, refines our understanding of mosquito fish genetics in their native range, evaluates important genotype–phenotype relationships, and identifies a potential new impact of climate change.     

Phenotypic integration in the feeding system of the eastern diamondback rattlesnake (Crotalus adamanteus)

Selection can vary geographically across environments and temporally over the lifetime of an individual. Unlike geographic contexts, where different selective regimes can act on different alleles, age‐specific selection is constrained to act on the same genome by altering age‐specific expression. Snake venoms are exceptional traits for studying ontogeny because toxin expression variation directly changes the phenotype; relative amounts of venom components determine, in part, venom efficacy. Phenotypic integration is the dependent relationship between different traits that collectively produce a complex phenotype and, in venomous snakes, may include traits as diverse as venom, head shape and fang length. We examined the feeding system of the eastern diamondback rattlesnake (Crotalus adamanteus) across environments and over the lifetime of individuals and used a genotype–phenotype map approach, protein expression data and morphological data to demonstrate that: (i) ontogenetic effects explained more of the variation in toxin expression variation than geographic effects, (ii) both juveniles and adults varied geographically, (iii) toxin expression variation was a result of directional selection and (iv) different venom phenotypes covaried with morphological traits also associated with feeding in temporal (ontogenetic) and geographic (functional) contexts. These data are the first to demonstrate, to our knowledge, phenotypic integration between multiple morphological characters and a biochemical phenotype across populations and age classes. We identified copy number variation as the mechanism driving the difference in the venom phenotype associated with these morphological differences, and the parallel mitochondrial, venom and morphological divergence between northern and southern clades suggests that each clade may warrant classification as a separate evolutionarily significant unit.     

Morphological and molecular diversity among populations of Quercus brantii Lindl. in western forest of Iran

Abstract

This study represents a preliminary step toward understanding the genetic structure of Persian oak in Iran. The genetic variability of Quercus brantii in Western forest of Iran was evaluated by amplified fragment length polymorphism (AFLP), chloroplast microsatellite and leaf morphology. Fifty-five trees from eight regions were sampled from across the range of Chaharmahal va Bakhtiari province of Iran. Twenty morphological traits were analyzed through clustering and ordination method. At morphological level, the applied statistics suggest that macromorphological traits significantly differentiate between populations. The overall sample shows a proportion of AFLP polymorphic markers of 92.1%, denoting a high level of variability. Based on AFLP data, differences among populations within geographic regions account for 11.6% of the total variation and only 0.57% is attributed to variation among regions. Based on chloroplast microsatellite (cpSSR), 34% of total variation was found among populations, suggesting a high within-population haplotype diversity. The dendrogram obtained from cpSSR showed a general pattern quite different from the pattern obtained from morphological analysis and AFLP markers.     

Morphological variability in Lophyra flexuosa (Fabricius, 1787) (Coleoptera,Cicindelidae) in desert countries is affected by sexual dimorphism and geographic aspect

Lophyra flexuosa, a eurytopic tiger beetle characterized by long phenological activity, wide geographic and altitudinal distribution, and occurring in the highest number of habitats among all Cicindelidae known from North Africa, was chosen to study its geographic variation in morphology and sexual dimorphism. Here, we found significant sexual dimorphism exhibited in larger body size of females and longer mandibles in males, which can be explained by different roles of particular sexes in courtship. Moreover, we recorded significant differences in body sizes between western and eastern Maghreb populations which could suggest genetic isolation between these populations. As the species is related to habitats placed close to the water reservoirs, which in the desert countries are under significant human pressure (including climate change), we expect a reduction of habitat occupied by this taxon. Therefore, the geographic morphological variability that we observe today in the tiger beetle Lophyra flexuosa in the future could lead to speciation.     

Untangling the Spurilla neapolitana (Delle Chiaje, 1841) species complex: a review of the genus Spurilla Bergh, 1864 (Mollusca: Nudibranchia: Aeolidiidae)

Spurilla neapolitana (Delle Chiaje, 1823) was considered to be a species with a broad geographic range and substantial colour variability; however, analyses of mitochondrial and nuclear gene data revealed that it is a complex of five distinct species. Further anatomical and morphological examinations determined that coloration is one of the main diagnostic traits for all five species, although some display substantial colour pattern variation. As a result of this study, S. neapolitana is determined to be restricted to the Mediterranean and eastern Atlantic. Spurilla sargassicola Bergh, 1871 from the Caribbean is redescribed and confirmed as a valid species. The name Spurilla braziliana MacFarland, 1909 is retained for western Atlantic and Pacific populations. Two new species are described herein. S purilla onubensis sp. nov. occurs in Europe, with a range overlapping that of S. neapolitana. Finally, S purilla dupontae sp. nov. is found in the Bahamas. © 2014 The Linnean Society of London     

Variation in complex mating signals in an “island” hybrid zone between Stenobothrus grasshopper species

Two grasshopper species Stenobothrus rubicundus and S. clavatus were previously shown to meet in a narrow hybrid zone on Mount Tomaros in northern Greece. The species are remarkable for their complex courtship songs accompanied by conspicuous movements of antennae and wings. We analyzed variations in forewing morphology, antenna shape, and courtship song across the hybrid zone using a geographic information system, and we documented three contact zones on Mount Tomaros. All male traits and female wings show abrupt transitions across the contact zones, suggesting that these traits are driven by selection rather than by drift. Male clines in antennae are displaced toward S. clavatus, whereas all clines in wings are displaced toward S. rubicundus. We explain cline discordance as depending on sexual selection via female choice. The high covariance between wings and antennae found in the centers of all contact zones results from high levels of linkage disequilibria among the underlying loci, which in turn more likely results from assortative mating than from selection against hybrids. The covariance is found to be higher in clavatus‐like than rubicundus‐like populations, which implies asymmetric assortative mating in parental‐like sites of the hybrid zone and a movement of the hybrid zone in favor of S. clavatus.     

Density‐dependent polyphenism and geographic variation in size among two populations of lubber grasshoppers (Romalea microptera)

1. Density‐dependent phase polyphenism occurs when changes in density during the juvenile stages result in a developmental shift from one phenotype to another. Density‐dependent phase polyphenism is common among locusts (Orthoptera: Acrididae). 2. Previously, we demonstrated a longitudinal geographic cline in adult body size (western populations = small adults; eastern populations = large adults) in the eastern lubber grasshopper (Romalea microptera) in south Florida. As lubbers are confamilial with locusts, we hypothesised that the longitudinal size cline was partly due to density‐dependent phase polyphenism. 3. We tested the effect of density, population, and density×population interaction on life‐history traits (pronotum length, mass, cumulative development time, growth rate) of, and proportion surviving to, each of the five instars and the adult stage in a 2 × 3 factorial laboratory experiment with two lubber populations, each reared from hatchling to adult at three different densities. 4. The effect of density on life history and survival was independent of the effects of population on life history and survival. Higher densities led to larger adult sizes (pronotum, mass) and lower survivorship. The western population had smaller adult masses, fewer cumulative days to the adult stage, and higher survivorship than the eastern population. 5. Our data suggest that lubber grasshoppers exhibit density‐dependent phase polyphenism initiated by the physical presence of conspecifics. However, the plastic response of adult size to density observed in the laboratory is not consistent with the relationship between phenotypes and adult density in the field. Genetic differences between populations observed in the laboratory could contribute to size and life‐history differences among lubber populations in the field.     

Morphological variation in the Chamaecytisus proliferus (L.f.) Link complex (Fabaceae: Genisteae) in the Canary Islands

FRANCISCO-ORTEGA, J., JACKSON, M. T., SANTOS-GUERRA, A. & FORD-LLOYD, B. V., 1993. Morphological variation in the Chamaecytisus proliferus (L.f.) Link complex (Fabaceae: Genisteae) in the Canary Islands . A multivariate study (Principal Component Analysis and Cluster Analysis, Warďs method) of 47 morphological traits from 164 populations of Chamaecytisus proliferus (L.f.) Link from the Canary Islands confirmed that this species complex is formed by seven morphological types. At least eight traits discriminated between these types. Patterns of variation follow a cline within Gran Canaria, Tenerife and La Palma. These results also show that morphological variation is greater in the eastern islands (i.e. Gran Canaria and Tenerife) than in the western islands (La Gomera, El Hierro and La Palma) and that no morphological differences are found between plants of typical tagasaste from wild and cultivated populations.