Geographic variation in Pemphigus populitransversus (Insecta: Aphididae)*†

The geographic variation of 33 morphological characters of two morphs of the gall-forming aphid Pemphigus populitransversus is studied in 214 locality samples. Among-locality variation ranges from 1 to 69% in the elongate morph and from 0 to 44% in the globular morph. The design of the study permits separation of interlocality correlations from intralocality correlations. The former are partly a function of the latter, confirming early observations on another Pemphigus species and on ticks. Factor analyses of both correlation matrices for both morphs yield four factors. Within localities these factors agree for the two morphs; among localities only one factor corresponds. Multiple discriminant analyses among localities of the two data sets do not correspond. There is little correlation between characters of stem mother and alate morphotypes within localities but such correlations are strong among localities. Maps are furnished for characters representing the independent dimensions of variation for each morph. Patterns of variation are shown to be significant by spatial autocorrelation analysis for both morphs but are much more marked in the elongate morph. Significant positive autocorrelation occurs up to 1000 km in the elongate morph – mostly only up to 200 km in the globular morph. There are two to three geographic variation patterns in the elongate morph, whereas in the globular morph the classes of patterns are less well defined and involve few characters. The environmental factors to which the globular and elongate morphs are adapting would appear to have different autocorrelation patterns. In each morph the patterns are clearly different and cannot be explained by a single microevolutionary process. The findings are compared with an earlier study in the related and largely sympatric P. populicaulis.     

Evolutionary relationships of Pemphigus and allied genera (Hemiptera: Aphididae: Eriosomatinae) and their primary endosymbiont,Buchnera aphidicola

Aphids harbor primary endosymbionts, Buchnera aphidicola, in specialized cells within their body cavities. Aphids and Buchnera have strict mutualistic relationships in nutrition exchange. This ancient association has received much attention from researchers who are interested in endosymbiotic evolution. Previous studies have found parallel phylogenetic relationships between non‐galling aphids and Buchnera at lower taxonomic levels (genus, species). To understand whether relatively isolated habitats such as galls have effect on the parallel relationships between aphids and Buchnera, the present paper investigated the phylogenetic relationships of gall aphids from Pemphigus and allied genera, which induce pseudo‐galls or galls on Populus spp. (poplar) and Buchnera. The molecular phylogenies inferred from three aphid genes (COI, COII and EF‐1α) and two Buchnera genes (gnd, 16S rRNA gene) indicated significant congruence between aphids and Buchnera at generic as well as interspecific levels. Interestingly, both aphid and Buchnera phylogenies supported three main clades corresponding to the galling locations of aphids, namely leaf, the joint of leaf blade and petiole, and branch of the host plant. The results suggest phylogenetic conservatism of gall characters, which indicates gall characters are more strongly affected by aphid phylogeny, rather than host plants.     

An Agrobacterium tumefaciens transformation protocol effective on a variety of cottonwood hybrids (genus Populus)

 We describe a protocol for Agrobacterium tumefaciens-mediated transformation of hybrid cottonwoods (Populus sections Tacamahaca Spach. and Aigeiros Duby). The protocol has allowed routine transformation of several economically important cottonwood hybrids (Populus trichocarpa Torr. & Gray×P. deltoides Bartr. ex. Marsh. and P. deltoides×P. nigra L.) that were previously difficult to transform. The procedure was applied to 11 different hybrid cottonwood genotypes and one P. deltoides genotype using kanamycin as the selection agent. Additional experiments showed a very strong interaction between auxin preculture and the effectiveness of various cytokinins for induction of shoot organogenesis. The data also demonstrated the superiority of Agrobacterium strain EHA105 over C58 and LBA4404 for T-DNA transfer based on transient assays with a reporter gene. Received: 16 June 1998 / Revision received: 5 February 1999 / Accepted: 14 April 1999     

Systematic status of the large squirrels (subgenus urosciurus) of the western Amazon basin

Patterns of geographic variation within and between regional samples of the large rufous western Amazon squirrels (Sciurus) are described based on univariate and multivariate analyses. Two species are clearly recognizable by independent patterns of geographic variation in measured characters, non‐overlapping qualitative cranial features, and certain pelage color characteristics. These species are a long‐muzzled, narrow‐skulled 5. spadiceus Olfers and a more normal‐muzzled, broader‐skulled S. igniventris Wagner. The two are sympatric over all of eastern Ecuador and most of Perú; only spadiceus is known from Bolivia.     

Comparative morphometrics by factor analysis in two species of diptera

Summary Covariational patterns of characters were compared in adults of two species of flies (Drosophila melanogaster and Musca domestica). Fourteen characters, homologous in the two organisms, were measured for both sexes in several laboratory strains of both species. Multiple factor analysis with rotation to an oblique simple structure was used to resolve the covariation into simpler patterns. Factor analyses were based on both the pooled within-group covariation as well as on total variation in each species. The five factors extracted from the correlation matrices of both species were quite similar after rotation (this was especially true for the within-group covariation). The following factors could be identified: a wing and a leg size factor, intercorrelated in the two species; a wing and thorax size factor; and separate factors affecting numbers of aristal and sternal setae. The present study is an example of the use of multiple factor analysis in comparative morphometricsContribution No. 36 from the Program in Ecology and Evolution at the State University of New York at Stony Brook. This study was initiated at the Department of Entomology of the University of Kansas. It was completed with support from grant GB-12587 of the National Science Foundation.     

Avian Response to Bottomland Hardwood Reforestation: The First 10 Years

Bottomland hardwood forests were planted on agricultural fields in Mississippi and Louisiana predominantly using either Quercus species (oaks) or Populus deltoides (eastern cottonwood). We assessed avian colonization of these reforested sites between 2 and 10 years after planting. Rapid vertical growth of cottonwoods (circa 2–3 m/year) resulted in sites with forest structure that supported greater species richness of breeding birds, increased Shannon diversity indices, and supported greater territory densities than on sites planted with slower‐growing oak species. Grassland birds (Spiza americana[Dickcissel] and Sturnella magna[Eastern Meadowlark]) were indicative of species breeding on oak‐dominated reforestation no more than 10 years old. Agelaius phoeniceus (Red‐winged Blackbird) and Colinus virginianus (Northern Bobwhite) characterized cottonwood reforestation no more than 4 years old, whereas 14 species of shrub‐scrub birds (e.g., Passerina cyanea[Indigo Bunting]) and early‐successional forest birds (e.g., Vireo gilvus[Warbling vireo]) typified cottonwood reforestation 5 to 9 years after planting. Rates of daily nest survival did not differ between reforestation strategies. Nest parasitism increased markedly in older cottonwood stands but was overwhelmed by predation as a cause of nest failure. Based on Partners in Flight prioritization scores and territory densities, the value of cottonwood reforestation for avian conservation was significantly greater than that of oak reforestation during their first 10 years. Because of benefits conferred on breeding birds, we recommend reforestation of bottomland hardwoods should include a high proportion of fast‐growing early successional species such as cottonwood.     

Trends in seedling growth and carbon‐use efficiency vary among broadleaf tree species along a latitudinal transect in eastern North America

Factors constraining the geographic ranges of broadleaf tree species in eastern North America were examined in common gardens along a ~1500 km latitudinal transect travers in grange boundaries of four target species: trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) to the north vs. eastern cottonwood (Populus deltoides) and sweet gum (Liquidambar styraciflua) to the south. In 2006 and 2007, carbon‐use efficiency (CUE), the proportion of assimilated carbon retained in biomass, was estimated for seedlings of the four species as the quotient of relative growth rate (RGR) and photosynthesis per unit tree mass (A_tree). In aspen and birch, CUE and RGR declined significantly with increasing growth temperature, which spanned 9 °C across gardens and years. The 37% (relative) CUE decrease from coolest to warmest garden correlated with increases in leaf nighttime respiration (R_leaf) and the ratio of R_leaf to leaf photosynthesis (R_%A). For cottonwood and sweet gum, however, similar increases in R_leaf and R_%A accompanied modest CUE declines, implying that processes other than R_leaf were responsible for species differences in CUE's temperature response. Our findings illustrate marked taxonomic variation, at least among young trees, in the thermal sensitivity of CUE, and point to potentially negative consequences of climate warming for the carbon balance, competitive ability, and persistence of two foundation species in northern temperate and boreal forests.     

PEMPHIGUS REVISITED: CHANGES IN GEOGRAPHIC VARIATION BUT CONSTANCY IN VARIABILITY AND COVARIATION

Samples of the gall-forming aphids Pemphigus populicaulis and P. populitransversus (both elongate and globular morphs) were re-collected at sites in eastern North America after 13 to 16 years. Twenty-three morphometric characters of the galls, stem mothers, and alate fundatrigeniae were analyzed by univariate and multivariate methods. Varying proportions of the variance of each character are attributable to the four levels of variation—locality, year, year by locality interaction, and among galls (within one year and locality). The year by locality interaction level generally has the greatest variation and is highly significant. Year and locality effects tend to be lower and not significant. The variance components do not exhibit trends with time. Geographic variation patterns of single variables or factor scores in the original and revisited populations show significant spatial structure overall but lack clear-cut spatial patterns, especially clines. Observable patterns of variation match results of the spatial analyses: most characters lack clear trends; patterns in the revisited data do not resemble patterns for the same variables in the original data. Variability profiles for characters change little over the time span and are comparable among and within localities. Covariation among characters over localities is largely maintained during the time interval despite the changes in patterns. Fluctuating interclonal competition among aphids on secondary hosts is believed to cause the marked heterogeneity in space and time among the aphids in the galls.     

Successful hybridisation of normally incompatible hybrid aspen (Populus tremula?��?P. tremuloides) and eastern cottonwood (P. deltoides)

Hybrid aspen (Populus tremula × P. tremuloides) belong to the section Populus. Eastern cottonwood (P. deltoides) is a member of the section Aigeiros within the genus Populus. These poplar sections are generally considered to be incompatible. Here, we describe successful hybridisation between these parents, producing an offspring family with 27 individuals. The hybrid character of individuals was proven by genotypes at 16 nuclear microsatellite loci. One individual was suspected to have more than the diploid chromosome number of 2n = 38 due to the observation of more than two alleles at several loci. This individual is a triploid, ascertained by flow cytometry. Two distinct growth classes of tall and dwarf plants were observed in the progeny, reflecting different degrees of postzygotic incompatibility. Two loci linked to the tested microsatellites have an effect on height growth. Some fast-growing individuals were micropropagated to test them for biomass performance together with other clones in field trials.     

Multilocus analysis of nucleotide variation and speciation in three closely related Populus (Salicaceae) species

Historical tectonism and climate oscillations can isolate and contract the geographical distributions of many plant species, and they are even known to trigger species divergence and ultimately speciation. Here, we estimated the nucleotide variation and speciation in three closely related Populus species, Populus tremuloides, P. tremula and P. davidiana, distributed in North America and Eurasia. We analysed the sequence variation in six single‐copy nuclear loci and three chloroplast (cpDNA) fragments in 497 individuals sampled from 33 populations of these three species across their geographic distributions. These three Populus species harboured relatively high levels of nucleotide diversity and showed high levels of nucleotide differentiation. Phylogenetic analysis revealed that P. tremuloides diverged earlier than the other two species. The cpDNA haplotype network result clearly illustrated the dispersal route from North America to eastern Asia and then into Europe. Molecular dating results confirmed that the divergence of these three species coincided with the sundering of the Bering land bridge in the late Miocene and a rapid uplift of the Qinghai‐Tibetan Plateau around the Miocene/Pliocene boundary. Vicariance‐driven successful allopatric speciation resulting from historical tectonism and climate oscillations most likely played roles in the formation of the disjunct distributions and divergence of these three Populus species.     

Characterization of three endophytic, indole-3-acetic acid-producing yeasts occurring in Populus trees

Three endophytic yeast, one isolated from stems of wild cottonwood (Populus trichocarpa), two from stems of hybrid poplar (P. trichocarpa × Populus deltoides), were characterized by analyzing three ribosomal genes, the small subunit (18S), internal transcribed spacer (ITS), and D1/D2 region of the large subunit (26S). Phenotypic characteristics of the yeast isolates were also obtained using a commercial yeast identification kit and used for assisting the species identification. The isolate from wild cottonwood was identified to be closest to species Rhodotorula graminis. The two isolates from hybrid poplar were identified to be species Rhodotorula mucilaginosa. In addition, the three yeast isolates were observed to be able to produce indole-3-acetic acid (IAA), a phytohormone which can promote plant growth, when incubated with l-tryptophan. To our knowledge, the yeast strains presented in this study were the first endophytic yeast strains isolated from species of Populus.     

Transpirational Water Loss in Invaded and Restored Semiarid Riparian Forests

The invasive tree, Tamarix sp., was introduced to the United States in the 1800s to stabilize stream banks. The riparian ecosystem adjacent to the middle Rio Grande River in central New Mexico consists of mature cottonwood (Populus fremontii) gallery forests with a dense Tamarix understory. We hypothesized that Populus would compensate for reduced competition by increasing its water consumption in restored riparian plots following selective Tamarix removal, resulting in similar transpiration (T) among stands. The northern study site included a Populus stand invaded by Tamarix (INV_N) and a restored Populus‐only stand (RES_N), as did a southern site (INV_S and RES_S) approximately 80 miles south. At each site, 20 × 20–m plots were established where up to 16 stems were monitored throughout the 2004 growing season using thermal dissipation sapflow sensors. Populus sapflux rates were greater in restored stands, suggesting those trees compensated for understory removal by using more water. Sapflow was scaled to estimate stand‐level T based on a quantitative assessment of sapwood basal area (A_sw) by species. Although exotic species represented 85 and 91% of the total stems in the invaded stands, it amounted to only 3% (INV_S) and 4% (INV_N) of the total A_sw, contributing proportionately less to T compared to Populus. Our results indicate that removing Tamarix from the Populus understory in this riparian forest had a minimal impact on stand water balance. Riparian restoration of the type discussed herein should focus primarily on enhancing riparian health rather than generating water.     

DISSECTING CORRELATED CHARACTERS: ADAPTIVE ASPECTS OF PHENOTYPIC COVARIATION IN MELANIZATION PATTERN OF PIERIS BUTTERFLIES

In this study we address the question of how much of the covariation among phenotypic characters observed in natural populations is adaptive. We examine covariation among a set of phenotypic characters that describe the wing-melanization pattern of Pieris butterflies. Previous functional analyses of thermoregulatory performance allow us to predict a priori whether and how different wing melanic characters should be correlated. We quantify and analyze the variation in the wing-melanization pattern within species for a series of Pieris populations from relatively cool environments in North America and compare these results with the predictions based on our adaptive hypothesis. We consider adaptive covariation both for biogeographic variation among populations and for seasonal polyphenism (phenotypic plasticity) within populations. Our hypothesis correctly predicts many of the qualitative features of covariation in melanization among major regions of the wings, at the level of biogeographic variation among populations, for both males and females of Pieris occidentalis. When within-population variation is considered, agreement with the adaptive predictions varies considerably in different populations for both P. occidentalis and P. napi males and females. Agreement for P. napi, particularly the females, is generally poorer than for P. occidentalis. In both species, there is a consistent difference in melanization pattern between alpine and arctic sites; this difference is discussed in relation to the differences in the radiative environment between these two types of “cold” habitats. Our results suggest that some important aspects of phenotypic correlation among wing melanic characters in Pieris are adaptive. We emphasize the important distinction between covariation and co-occurrence of characters, and we discuss these results in relation to the extensive biogeographic variation and phenotypic plasticity (seasonal polyphenism) in Pieris wing-melanization patterns.     

Complex clines: the predictivity of complicated patterns of geographic variation portrayed by multivariate analysis

The stability of complex patterns of geographic variation was investigated by assessing the congruence between multivariate ordinations derived from randomly chosen real characters. Two series of populations were analysed representing two situations with complex patterns of geographic variation. The first, a ‘Eurasian’ series of populations, showed a strongly structured hierarchical pattern, the second, an ‘eastern’ series of populations, showed a more subtle complex pattern of smooth clines and steps. The characters were selected from a total of 81 (Eurasian) or 61 (eastern) within-population independent characters from six different systems. The congruence between ordinations of the geographical populations was measured by the rotational fit statistic, R². Three procedures were used to compare ordinations based on from two to up to 80 characters randomly chosen to give: A, completely independent character sets; B, subsets compared to the total set; and C, potentially overlapping sets. All three procedures showed that congruence between the ordinations was asymptotic in relation to character number. This relationship was described by one of two mathematical models (procedure B did not result in a hyperbolic model as found with simple patterns of geographic variation). Generally speaking, once a sufficient number of characters are used, the complex patterns of geographic variation are stable, reliable and predictive and not substantially influenced by character choice. The strongly structured hierarchical pattern required 15 or so characters to achieve reliability whilst the more subtle patterns required 20 or so characters. However, the addition of further characters does improve reliability in both cases. The greater percentage of variance portrayed by three-dimensional ordinations compared to two-dimensional ordinations is achieved at the cost of lower congruence when a sufficient number of characters are used. If case studies of geographic variation were to adopt these procedures (preferably using completely independent character sets; procedure A) the reliability of their results would be indicated).     

Avian Density and Nest Survival on the San Pedro River: Importance of Vegetation Type and Hydrologic Regime

Abstract: Lowland riparian vegetation in the southwestern United States is critically important for maintaining a high richness and density of breeding birds. Further investigation is needed within riparian corridors, however, to evaluate the relative importance of vegetation type and hydrologic regime for avian density and nest survival as targets for regional conservation or restoration efforts. We estimated the densities of 40 bird species and for species grouped on the basis of nest height and dependence on surface water in gallery cottonwood–willow (Populus spp.–Salix spp.) forests, saltcedar (Tamarix spp.) shrub lands, and terrace vegetation types along a gradient in the hydrologic regime of the San Pedro River, Arizona, USA. We also assessed nest survival for shrub-nesting insectivores and herbivores. Canopy-nesting birds as a group and 14 individual bird species reached their greatest densities in cottonwood forests regardless of the hydrologic regime. Water-dependent birds as a group reached their highest density in both intermittent- and perennial-flow cottonwood stands, but certain species occurred almost exclusively in perennial-flow sites. Two shrub-nesting species and the brown-headed cowbird (Molothrus ater) were most abundant in saltcedar shrub lands, and the brown-headed cowbird was most abundant in saltcedar stands with intermittent flows. Mesquite (Prosopis spp.) and big sacaton (Sporobolus wrightii) grassland each maintained the highest densities of certain species within ≥1 hydrologic regime. Shrub-nesting insectivores had the greatest nest survival in cottonwood, including Arizona Bell's vireo (Vireo bellii arizonae), and also had lower proportions of nests parasitized and preyed upon, although 95% confidence intervals among vegetation types overlapped. Nest survival for both shrub-nesting insectivores and herbivores was lowest in intermittent-flow saltcedar, although, again, confidence intervals overlapped. Nest survival was lower in parasitized than nonparasitized nests in mesquite and across vegetation types for Arizona Bell's vireo and in cottonwood for Abert's towhee (Pipilo aberti). Riparian management that maintains heterogeneous riparian vegetation types, including floodplain vegetation comprising cottonwood–willow gallery riparian forests with some stretches of perennial flow, are important for maintaining the high diversity and abundance of breeding birds on the San Pedro River and probably across the region. Cottonwood stands also appear to maintain highest nest survival for some shrub-nesting birds.     

Expression of organomercurial lyase in eastern cottonwood enhances organomercury resistance

Summary Release of inorganic mercury pollutants into shallow aquatic environments has resulted in the bacterial production of a more toxic organic mercury species, methylmercury. The bacterial organomercurial lyase (MerB) catalyses the protonolysis of the carbon-mercury bond and releases Hg(II), a less toxic, non-biomagnified form of mercury. Our objective was to engineer eastern cottonwood (Populus deltoides), a fast-growing tree adapted to growth in riparian environments, with the merB gene to explore its potential for phytoremediation of mercury. We produced multiple eastern cottonwood clones expressing a modified bacterial merB gene, confirmed that the gene was expressed in the transclones and tested the regenerated plants for their ability to tolerate exposure to an organic mercury source, phenylmercuric acetate (PMA), in vitro and in hydroponic culture, compared to wild-type control trees. Transgenic merB plants expressed high levels of MerB protein and showed some evidence of higher resistance to the organic mercury than wild-type plants, producing longer roots under exposure to PMA in vitro, although hydroponic culture results were inconclusive. Our results indicate that in order for merB to be useful in eastern cottonwood trees designed to degrade methylmercury at mercury-contaminated aquatic sites, it will probably need to be combined with other genes such as merA.     

Aphid-ant interaction reduces chrysomelid herbivory in a cottonwood hybrid zone

In a cottonwood (Populus) hybrid zone, Chaitophorus aphids attract aphid-tending ants which subsequently reduce herbivory by the leaf-feeding beetle, Chrysomela confluens. Observations and experimental manipulations of aphids and beetle larvae on immature cottonwood trees demonstrated that: 1) via their recruitment of ants, aphids reduced numbers of beetle eggs and larvae on the host; 2) these interactions occurred within a few days of the host being colonized by aphids; and 3) although aphid colonies were ephemeral, their presence resulted in a 2-fold reduction in beetle herbivory. The aphid-ant interaction is most important in the hybrid zone where 93% of the beetle population is concentrated (for reasons unrelated to aphids and ants). Because beetle defoliation of immature trees is high (ca. 25%), the indirect effect of aphids in reducing herbivory is likely more beneficial to trees in the hybrid zone than in adjacent pure zones where beetle herbivory is virtually absent. Tree genotype likely affects the impact of the aphid-ant interaction on trees within the hybrid zone, since levels of herbivory differ between sympatric Fremont and hybrid cottonwoods.     

THE GEOGRAPHY OF MITOCHONDRIAL DNA VARIATION,POPULATION STRUCTURE,HYBRIDIZATION, AND SPECIES LIMITS IN THE FOX SPARROW (PASSERELLA ILIACA)

Geographic variation in mitochondrial DNA (mtDNA) restriction sites was studied in the fox sparrow (Passerella iliaca). Seventy-eight haplotypes were found. Haplotypes fall into four phylogeographic groups that correspond to groups defined by plumage characters. The geographic distribution of these four groups does not appear congruent with mtDNA patterns in other vertebrates. Within each group, there is little geographic variation in mtDNA restriction sites, although there is geographic variation in plumage coloration and body size. The evolution of mtDNA diversity in fox sparrows seems best explained by vicariant events rather than isolation by distance. The mtDNA evidence suggests that Passerella megarhyncha and Passerella schistacea, two nonsister taxa that occur in western North America, have independently undergone bottlenecks. Hybridization is limited between all pairs of taxa except P. megarhyncha and P. schistacea, where mtDNA evidence suggests a narrow contact zone along the interface of the Great Basin and Sierra Nevada/Cascades. Morphometric characters intergrade over a broader area, suggesting that different processes are responsible for the two gradients. The occurrence of limited backcrossing among taxa suggests that cytoplasmic-nuclear incompatibility is lacking. The number of biological species would range from one to four, depending on the degree of hybridization tolerated. The mtDNA and plumage characters suggest four phylogenetic species: P. iliaca, P. megarhyncha, P. unalaschcensis, and P. schistacea.     

A land snail's view of a fragmented landscape

Habitat fragmentation may influence the genetic structure of populations, especially of species with low mobility. So far, these effects have been mainly studied by surveying neutral markers, and much less by looking at ecologically relevant characters. Therefore, we aimed to explore eventual patterns of covariation between population structuring in neutral markers and variation in shell morphometrics in the forest‐associated snail Discus rotundatus in relation to habitat fragment characteristics. To this end, we screened shell morphometric variability and sequence variation in a fragment of the mitochondrial 16S rDNA gene in D. rotundatus from the fragmented landscape of the Lower Rhine Embayment, Germany. The 16S rDNA of D. rotundatus was highly variable, with a total of 118 haplotypes (384 individuals) forming four clades and one unresolved group. There was a geographic pattern in the distribution of the clades with the river Rhine apparently separating two groups. Yet, at the geographic scale considered, there was no obvious effect of fragmentation on shell morphometrics and 16S rDNA variation because G_ST often was as high within, as between forests. Instead, the age of the habitat and (re‐)afforestation events appeared to affect shell shape and 16S rDNA in terms of the number of clades per site. The ecologically relevant characters thus supported the presumably neutral mitochondrial DNA markers by indicating that populations of not strictly stenecious species may be (relatively) stable in fragments. However, afforestation after large clearcuts and habitat gain after the amendment of deforestation are accompanied by several, seemingly persistent peculiarities, such as altered genetic composition and shell characters (e.g. aperture size). © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 839–850.