Article

Morphometric correlates of karyotype and host plant in genus Euceraphis (Hemiptera: Aphididae)

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Abstract

Abstract Samples of birch- and alder-feeding aphids of genus Euceraphis were measured and analysed using multiple discriminant analysis (canonical variates) to find out if morphological variation correlated with previously reported differences of karyotype and host association. The dataset comprised groups of specimens defined solely by locality and collection date. Mean scores on the first two canonical variates clustered the samples fully in accordance with their karyotypes and host plants, confirming the existence of a number of morphologically similar but distinct host-specific taxa within the E. betulae group. Three new species are described: Euceraphis borealis Blackman, sp.n. on Betula glandulosa and B. nana, Euceraphis papyrifericola Blackman, sp.n. on B. papyrifera, and Euceraphis quednaui Blackman, sp.n. on B. occidentalis. A key is provided to alate viviparae of the genus.

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... (Blackman et al., 1977), Myzus persicae group (Blackman, 1987; Margaritopoulos et al., 2000), Aphis gossypii Glover (Margaritopoulos et al., 2006) and Euceraphis spp. (Blackman & De Boise, 2002). Previous reports in the literature suggest that H. pruni and H. amygdali differ from each other in the length-width ratio of the cornicle of adult apterae and the number of sensoria on the third antennal segment of alatae (Börner & Heinze, 1957; Eastop, 1966). ...
... This technique can effectively separate environmental from genetic components of variance and even distinguish between different genotypes, as has been demonstrated using aphid clones of closely related species reared under laboratory conditions (Blackman & Spence, 1994). In addition, the technique has proved effective when applied to samples from field populations (e.g. Blackman & De Boise, 2002; Margaritopoulos et al., 2006). The separation obtained in the present study might also reflect some environmental influences in addition to the – genetic differences between the three groups. ...
Article
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Morphometric variation of individuals in field collected samples of the Hyalopterus pruni complex from various Prunus species and regions of Greece was examined, to determine whether this variation is correlated with the host-trees from which the aphids originated. Morphometric data for 13 parameters of aphids from 74 field samples (760 adult apterae) were analysed by canonical variates analysis (CVA). Each sample was collected from a different tree and consisted of individuals from 2-3 neighbouring leaves from the same branch. Each field sample was used as a grouping factor in the CVA. The analysis produced three clusters corresponding to the taxa indicated by previous studies using allozyme markers (i.e., Hyalopterus pruni on apricot, black-thorn, plum and cherry, H. amygdali A on almond and H. amygdali B on peach). The separation was independent on the geographical origin of the samples. However, host associations within the complex were not absolute, and in particular the samples from apricot appeared to include both H. pruni and H. amygdali A. In contrast to previous views, the present study showed that the taxa have their own distinct morphology. Lastly, discriminant functions for separating the adult apterae of the taxa are given.
... In particular, morphological separations or affinities were confirmed through comparisons between the species-groups. Multivariate morphometric analysis is generally used to represent complex, multidimensional patterns of variation between host-related populations in polyphagous aphids [20,21,22] and has also been proven to be a powerful and reliable method for separating closely-related aphid taxa [23,24,25]. However, in contrast to previous studies, we employed morphometric analysis to cluster species and confirm the relationships between the previously classified species-groups in Aphis. ...
... To obtain the mean value of each character, we examined a total of 1,264 slide mounted specimens from 40 species (Table S2). The continuous characters measured in this study have been shown to be useful in other morphometric studies of aphids [22,23,25,51], while the meristic characters were new characters adopted from morphometric and biometric analyses of Aphis [52,53,54]. Length measurements were taken using Image Lab ver. ...
Article
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The species-rich genus Aphis consists of more than 500 species, many of them host-specific on a wide range of plants, yet very similar in general appearance due to convergence toward particular morphological types. Most species have been historically clustered into four main phenotypic groups (gossypii, craccivora, fabae, and spiraecola groups). To confirm the morphological hypotheses between these groups and to examine the characteristics that determine them, multivariate morphometric analyses were performed using 28 characters measured/counted from 40 species. To infer whether the morphological relationships are correlated with the genetic relationships, we compared the morphometric dataset with a phylogeny reconstructed from the combined dataset of three mtDNA and one nuclear DNA regions. Based on a comparison of morphological and molecular datasets, we confirmed morphological reduction or regression in the gossypii group unlike in related groups. Most morphological characteristics of the gossypii group were less variable than for the other groups. Due to these, the gossypii group could be morphologically well separated from the craccivora, fabae, and spiraecola groups. In addition, the correlation of the rates of evolution between morphological and DNA datasets was highly significant in their diversification. The morphological separation between the gossypii group and the other species-groups are congruent with their phylogenetic relationships. Analysis of trait evolution revealed that the morphological traits found to be significant based on the morphometric analyses were confidently correlated with the phylogeny. The dominant patterns of trait evolution resulting in increased rates of short branches and temporally later evolution are likely suitable for the modality of Aphis speciation because they have adapted species-specifically, rapidly, and more recently on many different host plants.
... distinguish even between different genotypes. It can also be usefully applied to samples from field populations (Blackman & De Boise, 2002; see also the review by Blackman, 1992 for the properties of the method). A criticism (e.g. ...
... (Blackman et al., 1977), Rhopalosiphum maidis Brown & Blackman (1988), Myzus persicae (Blackman, 1987;Margaritopoulos et al., 2000;Margaritopoulos et al., 2005) and Euceraphis spp. (Blackman & De Boise, 2002). However, it is unusual to find specificity at the level of the host family within what is regarded as one aphid species. ...
Article
Aphis gossypii Glover is a polyphagous aphid pest with a worldwide distribution. However, there is evidence that on a global scale the name A. gossypii is being applied to a number of forms with different life cycles and/or host-plant associations. Morphometric variation of A. gossypii samples from crops and non-cultivated plants in many parts of the world was examined, to determine whether this variation is correlated with the hosts from which the aphids originated. Samples of A. gossypii were collected from Cucurbitaceae and Malvaceae in Europe, and from Compositae in various parts of the world. Morphometric data for 13 parameters measured from 97 clonal lineages (728 specimens) and 27 field-collected samples (313 specimens) were analysed by a series of canonical variates analyses, using the field sample/clonal lineage as grouping factor. Clonal lineages were reared on a common host in controlled conditions to standardize the effect of host and environment on morphology. The analyses provided a clear morphometric separation of the aphids originating from Compositae and those collected on Cucurbitaceae and Malvaceae, regardless of the geographical origin of the aphids and the host plant on which they were reared. This indicates that within A. gossypii there are two widely distributed host races or subspecies with different plant family associations. The taxonomic implications are discussed.
... Comments. these aphids are not identifiable with the existing keys to the genus Euceraphis (Blackman & de Boise, 2002;Blackman & eastop, 2016): several features are intermediate, falling between the values given in the keys' couplets. the small number of specimens prevents deciding whether these intermediate characteristics represent intraspecific variation or the collected specimens belong to a previously unknown taxon. ...
Article
The first study of the aphid fauna from Nenets Autonomous okrug has been made; 64 species are reported from this territory, the most north-eastern of Europe. Two forms with unclear taxonomic status are described: Acyrthosiphon sp. aff. knechteli, a polyphagous species found on plants in six different families; and Pleotrichophorus sp. aff. glandulosus on Tanacetum bipinnatum (L.) Sch. Bip. Previously considered a synonym of Aulacorthum rufum Hille Ris Lambers, 1947, A. pirolacearum Szelegiewicz, 1967, is re-established.
... Comments. Specimens from Chukotka differ slightly in morphology from those described by Blackman and De Boise (2002). Our specimens have a slightly shorter base of the 6th antennal segment-306-372 (320-390 in ...
Article
The first study of the aphid fauna from Chukotka Autonomous Okrug has been made; 18 species are reported from this territory, the most north-eastern of Russia and of the Palaeartic Region. Five new species are described: Aphis aquilonalis sp. nov. on Rumex arcticus Trautv.; Aphis beringiensis sp. nov. were collected by yellow trap; Macrosiphoniella (Asterobium) jaroslavi sp. nov. on Artemisia arctica Less. and Artemisia tilesii Ledeb.; Macrosiphum chukotense sp. nov., a polyphagous species found on plants in nine different families; and Pleotrichophorus tuberculatus sp. nov. on Artemisia tilesii Ledeb. In addition, the male and oviparous female of Acyrthosiphon boreale Hille Ris Lambers, 1952 and the fundatrix and male of Pleotrichophorus knowltoni Corpuz-Raros and Cook, 1974 are described for the first time.
... The results of the PCA evidently confirm morphological differentiation of studied genera (Fig. 7); the characters contributing most to morphometric discrimination are the lack of cauda, PT/BASE as well as the shape of frons and siphunculi. The PCA is a valuable method to verify morphological separation of aphid morphs (Depa & Kanturski 2014) as well as species and genera (Tizado & Nieto Nafría 1994;Blackman & De Boise 2002;Ortiz-Rivas et al. 2009;Mier Durante et al. 2013). Our analysis has confirmed that differences between A. puchovi and B. eastopi are considerable and that representatives of both taxa should be treated as species from separate genera. ...
Article
The aphid genus Blackmania gen. nov. is described. In the genus, B. eastopi sp. nov. associated with Polygonum equisetiforme (Polygonaceae) from Israel and Cyprus is described and illustrated. Morphologically , the new genus is similar to the genus Acaudella in respect to the lack of a cauda. Acaudella puchovi, associated with Atraphaxis caucasica (=A. buxifolia) and A. spinosa (Polygonaceae) from Uzbekistan and Israel, is re-described and the apterous viviparous female is figured for the first time. The lectotype and paralectotypes of A. puchovi are also designated. An identification key to known species of the tribe Macrosiphini without cauda is provided. The morphological separation of B. eastopi gen. nov., sp. nov. from A. puchovi is visualized using principal component analysis.
... Our analysis confirmed that differences between them are significant and that representatives of both taxa should be treated as separate species within this genus. It also confirms that mentioned methods are valuable both in verifying morphological separation of aphid morphs, species and genera (Blackman and De Boise, 2002;Depa and Kanturski, 2014;Kanturski and Wieczorek, 2015;Mier Durante et al., 2013;Ortiz-Rivas et al., 2009;Tizado and Nieto Nafria, 1994). As sexuales are known only in D. aceris aceris and D. fugans, their characteristics were not considered in the performed statistical analysis. ...
Article
The genus Drepanosiphoniella Davatchi, Hille Ris Lambers & Remaudière, 1957 of the subfamily Drepanosiphinae (Hemiptera: Aphididae) is revised using comparative morphology of parthenogenetic and sexual generations. On the basis of thorough examination of morphological characters, a full species status of D. fugans Remaudière & Leclant, 1972 stat. rev. is given, and supported by using statistical analysis – principal component analysis (PCA) and multiple discriminant analysis (canonical variates analysis). A new species, D. remaudièrei sp. n. Wieczorek, is described and figured in detail on the basis of specimens collected in Morocco. Fundatrix, oviparous female and apterous male of D. aceris aceris Davatchi, Hille Ris Lambers & Remaudière, 1957 are described and figured. Original keys to the identification of the known species and morphs of the genus Drepanosiphoniella as well as differences with related species are given. Notes about the distribution and host plants of the studied taxa are supplied. All species studied seem to be montane elements associated with Acer monspessulanum or its subspecies
... Multiple discriminant analysis, specifically canonical variate analysis (CVA), has proved to be a powerful and reliable method of separating closely-related aphid taxa (Blackman, 1992;Blackman and Spence, 1994;Blackman and De Boise, 2002), and effectively discriminates between M. persicae persicae and M. persicae nicotianae (Blackman, 1987;Margaritopoulos et al., 2000), but has not previously been applied to the genotypes studied in Japan. Here we have used this technique to examine whether the morphology of Japanese clones from tobacco is different from that of clones from other crops, and to compare the Japanese clones with Greek genotypes of M. p. persicae and M. p. nicotianae identified by previous morphometric studies (Margaritopoulos et al., 2000. ...
Article
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There have been independent long-term studies of tobacco-adapted populations of Myzus persicae in Japan and Greece, but until now no direct comparisons have been made between them. In Japan, as in Greece, tobacco aphids can reproduce sexually to some extent so that there is the potential for hybridisation with M. persicae from other plants on their common primary host, the peach. To determine whether Japanese tobacco aphids agree morphologically with those in Greece, and hence fit the concept of M. persicae ssp. nicotianae, multivariate morphometric analysis was applied to 16 clones of M. persicae s.lat. from various localities in Japan. The clones represented 14 different "colour-esterase forms" collected from tobacco and other crops over a 20-year period and kept in laboratory culture. The analysis discriminated most of the clones from tobacco from those collected on other crops. When Japanese samples were analysed together with those from Greece, the tobacco-feeding clones of both countries had a similar pattern of morphometric correlations, discriminating them from those on other crops, even though they were reared under different conditions. Thus, the genomic integrity of M. p. nicotianae seems to be preserved across a wide geographical scale and even in regions where interbreeding can occur. The possible origin and evolution of the tobacco-adapted subspecies are discussed.
... In a morphometric study, the geographic variation of 33 morphological characters of the gall-forming aphid, Pemphigus populicaulis, was investigated in Eastern North America (Sokal et al. 1980). Canonical variates analysis has been extensively used as a powerful tool to discriminate and resolve the taxonomic status among related insect species and closelyrelated aphid taxa (Rakauskas 1998, Sanmartin & Martin Piera 1999, Blackman & de Boise 2002, Williams & Langor 2002, Barbagallo & Cocuzza 2003, Margaritopoulos et al. 2006, Poulios et al. 2007. ...
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Multivariate morphometric studies are often carried out on different aphid taxa in order to analyze population variation and species discrimination in relation with different factors such as host plant, temperature and geography, which have significant effect on aphid morphology. The chrysanthemum aphid, Macrosiphoniella sanborni (Hemiptera: Aphididae), is a widespread pest on cultivated chrysanthemum throughout the world. In the present study 21 morphological characters in 11 populations of M. sanborni collected from several geographic locations in Iran were examined. Multivariate morphometric techniques, canonical variates analysis (CVA) and cluster analysis were used to determine whether these populations can be reliably morphologically discriminated. The results showed such discrimination between populations. The Kermanshah population (located in western Iran) clearly separated from other populations, indicating the presence of a morphologically distinct group. It could be concluded that presence of a geographic barrier and climatic conditions are among the most important factors which contribute to the differentiation between populations. Other complimentary studies such as genetic investigation will provide a better understanding of the evolutionary implications of the population structure.
... Canonical variates analysis has proved to be a powerful technique that helps to resolve taxonomic difficulties in closely related aphid taxa (Margaritopoulos et al. 2006, Poulios et al. 2007, Lozier et al. 2008). Aphid samples from field populations have been studied using this technique (Blackman & De Boise 2002, Poulios et al. 2007). Although factors such as environmental conditions and natural enemies may have considerable effect on aphid morphology (Dixon 1998, Margaritopoulos et al. 2000), it seems that the morphological separation in this study is mostly due to host-related differences because samples were collected from host plants in a small scale range with similar climatic conditions during about three weeks in the spring. ...
Data
Morphometric analyses were carried out on some populations of Aphis craccivora Koch (Hemiptera: Aphididae) associated with different host plants. Twelve morphological characters were measured. Two multivariate discriminant analyses, Canonical variates analysis and Cluster analysis were employed to discriminate individuals of A. craccivora populations on a morphological basis. Both Canonical variates analysis and Cluster analysis revealed significant differences among A. craccivora collected from different host plants. A stepwise Canonical variates analysis selected four morphological characters (siphunculus length, ultimate rostral segment length, basal diameter of siphunculus and processus terminalis length) from which two (siphunculus length and ultimate rostral segment length) showed the highest contribution to the separation of host-associated populations and 87.6% of individuals were reclassified correctly into their original populations. The result of morphometric analyses performed here revealed the existence of three morphologically differentiated groups within A. craccivora associated with different host plants which provide evidence for the presence of host related forms. This study demonstrates that morphometric analysis of morphological characters can discriminate among host related groups based on size variation and clearly separates the individuals of A. craccivora. The aphid population associated with Robinia pseudoacacia L. was clearly separated from other populations. The most important discriminatory characters of A. craccivora and their functions in relation to adaptation of populations with different host plant species are discussed. We concluded that A. craccivora populations are not homogeneous morphological entities and represent different host-associated forms. The results presented here for the morphological structure of A. craccivora provide the framework to investigate the genetic and biological differences among these entities so that we could be able to exactly evaluate their taxonomic situation.
... Multiple discriminant (canonical variates) analysis was performed using the 23 samples as the grouping variable . This method maximises the differences between groups (= samples), and has proved to be powerful in separating the environmental and genetic components of variance (Blackman & Spence 1994) and thus resolving differences between closely-related species (Blackman 1992, Blackman & De Boise 2002). Analysing the data for all 23 groups, the first five canonical variates (CVs) accounted for 89.3% of the total variance in the data, of which CV1 accounted for 46.2%, CV2 for 22.4%, CV3 for 11.3%, CV4 for 5.8% and CV5 for only 3.6%. ...
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Information is given concerning the karyotypes of 160 species of Japanese Aphididae, including new data for 130 species. Most karyotypes are illustrated by means of idiograms. The cytological and taxonomic significance of the data is discussed. Species with X1X2O sex determination have been found in Colopha, Colophina and Tetraneura (Pemphiginae) and in Stomaphis (Lachinae). Genera in which fusions or dissociations seem to have occurred frequently in the course of speciation include Kaltenbachiella, Tetraneura (Pemphiginae); Chaitophorus, Euceraphis, Symydobius (Drepanosiphinae); Amphorophora, Aulacorthum (subgen. Neomyzus), Cavariella, Melanaphis and Vesiculaphis (Aphidinae). The cytological work of Shinji is reviewed and some discrepancies with the present findings are discussed. © 1986, Japan Mendel Society, International Society of Cytology. All right reserved.
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Autosomal dissociations are a common feature of aphid karyotype evolution, but multiple X chromosome systems are rare. Birch-feeding aphids of the genus Euceraphis, however, have X1X2O males as a general rule, X1 being always much larger than X2. Only one species has XO males, and this condition appears to be secondary. Most Euceraphis karyotypes also have one or more, usually heterochromatic, elements that occur in the same numbers in both males and females, yet behave like X chromosomes at male and female meiosis I. They appear to be supernumerary, non-functional X chromosomes, although showing greater within-species stability in size and number than typical B chromosomes. Euceraphis gillettei forms a separate group within the genus and feeds on alders (Alnus species), yet has a similar system, and the two most closely related genera, Symydobius and Clethrobius, also have additional chromosomal elements possibly representing non-functional X chromosomes. Thus the multiple X chromosome system in these aphids seems to be a primitive condition.
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This paper deals primarily with new species of Aphididæ, on Artemisia, with notes from type slides of some described species on the same host, in the hope of clearing up some confusion and correcting typographical errors in literature. Three hitherto undescribed species on Populus, Valeriana and Pedicularis, respectively, have been added.
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Provide keys and descriptions of tree-dwelling aphids.
Article
Variation within and between samples of emigrant alatae from galls on identified Pistacia species was studied using bivariate and multivariate techniques. The Geoica utricularia complex seems to include at least five taxa, including two newly identified; G.utricularia Passerini sensu stricto, on Pistacia terebinthus; G.muticae Mordvilko, on P.mutica; G.rungsi Davatchi & Remaudière, on P.atlantica; G.harpazi sp.n., also on P.atlantica; and G. wertheimae sp.n. on P. palaestina. The emigrant alatae of the two new species are described, together with apterous exules reared from them, and a key is provided to emigrant alatae of all eight Geoica species now known to induce galls on Pistacia. The taxonomic status of anholocyclic, grass-feeding populations in the Geoica utricularia group is discussed.
An annotated list of Aphididae of North America
  • C.F. Smith
  • C.S. Parron
  • C.F. Smith
  • C.S. Parron
cordi-folia and B. populifolia, 2.vii.77 (Blackman)
  • New / Brunswick
  • Fredericton
9 al. viv. /, New Brunswick, Fredericton, ex B. cordi-folia and B. populifolia, 2.vii.77 (Blackman); 12 al. viv. /, New Brunswick, Fredericton, ex B. papyrifera, 30.vi-2.vii.77 (Blackman);
Aphids of the genus Euceraphis Walker 1870 (Homoptera, Aphidinea) in the Far East
  • M A Palmer
Palmer, M.A. (1952) Aphids of the Rocky Mountain Region. Thomas Say Foundation, Denver. Pashtshenko, N.F. (1984) Aphids of the genus Euceraphis Walker 1870 (Homoptera, Aphidinea) in the Far East. Systematics of the Insects from the Far East (ed. by P. A. Lehr), pp. 8±17. Akademie Nauk SSSR, Vladivostok. Pashtshenko, N.F. (1988) Suborder Aphidinea ± Aphids. Keys to Insects of the Far East of USSR, 2 (ed. by P. A. Lehr), pp. 546±686. `Nauka', Leningrad. Smith, C.F. & Parron, C.S. (1978) An annotated list of Aphididae of North America. Technical Bulletin of North Carolina Agricultural Experiment Station, 255, 1±428.
Vancouver, ex B. papyrifera, 13.vii
  • British / Columbia
2 al. viv. /, British Columbia, Vancouver, ex B. papyrifera, 13.vii.87 (Chan); 1 al. viv. / (sexupara), Manitoba, Clear Lake, ex B. papyrifera, 26.viii.64 (Robinson);
A new and little known species of Aphididae from Japan (Homoptera). KontyuÃ, 38, 252±256
  • M Sorin
Sorin, M. (1970) A new and little known species of Aphididae from Japan (Homoptera). KontyuÃ, 38, 252±256. Accepted 17 July 2001
Suborder Aphidinea ± Aphids. Keys to Insects of the Far East of USSR
  • N F Pashtshenko
Pashtshenko, N.F. (1988) Suborder Aphidinea ± Aphids. Keys to Insects of the Far East of USSR, 2 (ed. by P. A. Lehr), pp. 546±686.`Nauka', Leningrad.
  • C P Gillette
  • M A Palmer
Gillette, C.P. & Palmer, M.A. (1931) Aphididae of Colorado. Part I. Annals of the Entomological Society of America, 24, 827±934.
An annotated list of Aphididae of North America. Technical Bulletin of North Carolina Agricultural Experiment Station
  • C F Smith
  • C S Parron
Smith, C.F. & Parron, C.S. (1978) An annotated list of Aphididae of North America. Technical Bulletin of North Carolina Agricultural Experiment Station, 255, 1±428.
Knowlton); Utah, Anogsa, 27.v.57 (Knowlton). 1 ovip
  • Box Utah
  • Elder Canyon
Utah, Box Elder Canyon, 9.v.57 (Knowlton); Utah, Anogsa, 27.v.57 (Knowlton). 1 ovip. /, U.S.A.: Utah, Logan Canyon, 25.x.57 (Knowlton).
The use of ordination techniques to discriminate within pest aphid species complexes. Ordination in the Study of Morphology
  • R L Blackman
Blackman, R.L. (1992) The use of ordination techniques to discriminate within pest aphid species complexes. Ordination in the Study of Morphology, Evolution and Systematics (ed. by J. T. Sorensen and R. Foottit), pp. 261±275. Elsevier, Amsterdam. Blackman, R.L. & Eastop, V.F. (1994) Aphids on the World's Trees. CAB International, Wallingford, U.K.
Aphids of the genus Euceraphis Walker 1870 (Homoptera, Aphidinea) in the Far East
  • N F Pashtshenko
Pashtshenko, N.F. (1984) Aphids of the genus Euceraphis Walker 1870 (Homoptera, Aphidinea) in the Far East. Systematics of the Insects from the Far East (ed. by P. A. Lehr), pp. 8±17. Akademie Nauk SSSR, Vladivostok.
28 (Knowlton); Utah, Pine Valley Mountains, 11.vii.63 (Knowlton)
  • Blacksmith Utah
  • Canyon
Utah, Blacksmith Canyon, 23.vi.28 (Knowlton); Utah, Pine Valley Mountains, 11.vii.63 (Knowlton); Utah, Huntaville, 20.vi.57 (Knowlton); Utah, Ogden Canyon, 20.vi.57 (Knowlton);
Blackman) (BMNH). Paratypes, 11 al. viv. /, same sample no. and data as holotype Other material examined (all ex Betula occidentalis), 116 al: Idaho, Cub River Canyon, 25.vii
  • Material
  • Holotype
  • Usnm Bmnh
Material examined. Holotype, al. viv. /, embryos karyotyped as 2n ˆ 11, from sample no. NA37/RLB 1269, U.S.A.: Utah, Franklin Basin, Logan Canyon, ex Betula occidentalis, 13.vi.77 (Blackman) (BMNH). Paratypes, 11 al. viv. /, same sample no. and data as holotype (BMNH, USNM, CNC). Other material examined (all ex Betula occidentalis), 116 al. viv. /, U.S.A.: Idaho, Cub River Canyon, 25.vii.64 (Knowlton); Utah, Logan Canyon, 19.vi.57, 21.vi.63, 26.vii.64, 7.vi.66, 1.vi.73, 17.v.74, 3±21.vi.76, 12±13.vi.77 (Knowlton, Pepper, Stroyan, Blackman); Utah, Provo Canyon and American Fork Canyon, 11.vi.76 (Blackman);
Biological flora of the British Isles, Betula nana L. & Betula glandulosa Michx
  • de Groot
A new and little known species of Aphididae from Japan (Homoptera)
  • Sorin
Aphididae of Colorado. Part I
  • Gillette
An annotated list of Aphididae of North America
  • Smith