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A molecular biosystematic study on North American Solidago and related genera (Asteraceae: astereae) based on chloroplast DNA RFLP analysis [microform].
Abstract
Thesis (Ph. D.)--University of Waterloo, 1996. Includes bibliographical references.
... a number of studies have applied molecular data to the resolution of systematic problems in astereae (suh & simpson 1990;morgan & simpson 1992;lane et al. 1996;Zhang 1996;noyes & rieseberg 1999;Urbatsch et al. 2003;Beck et al. 2004), so a considerable body of molecular data has begun to accumulate for the tribe. other than a chloroplast dna restriction site analysis (Zhang 1996;semple et al. 1999), there has been no focus as yet for dna sequence analysis on solidago, although a few sequences are available from other work. ...
... a number of studies have applied molecular data to the resolution of systematic problems in astereae (suh & simpson 1990;morgan & simpson 1992;lane et al. 1996;Zhang 1996;noyes & rieseberg 1999;Urbatsch et al. 2003;Beck et al. 2004), so a considerable body of molecular data has begun to accumulate for the tribe. other than a chloroplast dna restriction site analysis (Zhang 1996;semple et al. 1999), there has been no focus as yet for dna sequence analysis on solidago, although a few sequences are available from other work. as part of an initial survey for suitable molecular markers, we analyzed its and ets sequence variation for several samples of solidago that represented the major subunits of the genus, as delimited by semple and cook (2006). ...
... a notable aspect of the its and ets sequence data was the striking lack of divergence among members of solidago. By contrast, its divergence in eupatorium l. another genus of asteraceae with a similar geographic range and occurrence in old field habitats, is 2-6% between species (12-43 bp differences; schmidt & schilling 2000; schilling et al. 2007); eupatorium is further differentiated from eutrochium raf., which are widely treated as congeners, by two indels (loss of a 13 bp region in eupatorium and addition of a 3 bp region in eutrochium) in addition to a minimum of 8% (33-61 bp) divergence. it should be noted that a previous chloroplast rflp analysis (Zhang 1996) reports a higher level of variation within solidago, and the resulting phylogeny strongly supports certain previously hypothesized groups. these contrasting levels of phylogenetic signal are best explained by the relative amount of dna sequence scored in each study. ...
Analysis of nuclear nbosomal ITS and ETS sequence data was used to assess the relationships of the garden ornamental Solidaster cv Lemore, a cultivar that was at one time believed to be an intergeneric hybrid (Solidago × Aster). Part of the analysis involved assessment of the generic placement of one of its putative parents, the Upland White Aster, Solidago ptarmicoides Because of its superficial external appearance this species is still often treated as an Aster, although there is evidence from several sources that suggests it should be classified within Solidago. Phylogenetic analysis of the combined ITS and ETS sequence data showed that there was strong support for the inclusion of S. ptarmicoides within Solidago, based on its placement in a well supported clade composed of all other sampled species of the genus. The ITS sequence data of Solidaster showed evidence of a hybrid origin based on the presence of several intraindividual base pair polymorphisms. Cloning experiments recovered two different individual ITS sequences, one identical to that obtained from S. ptarmicoides and a second that matched sequences obtained for S canadensis Thus DNA sequence data suggested that Sohdaster cv. Lemore is a hybrid goldenrod that involved a cross between S. ptarmicoides and S. canadensis The data further indicated that another possible candidate, Euthamia graminifolia, is not a parent of Solidaster. Also notable was the striking lack of sequence divergence for ITS and ETS among species of Solidago, suggesting that estimation of phylogenetic relationships within the genus will require more rapidly evolving markers.
... Another cpDNA restriction site study (Morgan & Simpson, 1992) identified a well (96% bootstrap) supported clade consisting of proposed Solidagininae genera (Solidago, Oreochrysum, and Tonestus). In contrast, proposed Solidagininae genera formed polyphyletic groups in analyses of another cpDNA restriction site dataset (Zhang, 1996) as well as ones based on nuclear ITS sequence data (Noyes & Rieseberg, 1999;Brouillet & al., 2001;Semple & al., 2002). These studies should be interpreted with caution, however, because none included sufficient sampling for a rigorous phylogenetic assessment of Solidagininae. ...
... Solidago (ca. 100 spp.) has been variously defined, with disagreement over the status of several possible segregate species or groups of species (reviewed in Zhang, 1996). Our minimal sampling of Solidago s.s. ...
... Further expansion of Solidago may still be warranted as future analyses may identify a clade composed of Solidago s.s., Oligoneuron, Chrysoma, the rayless Solidago discoidea, and Oreochrysum parryi. The cpDNA data of Zhang (1996) strongly supported a clade including Solidago s.s., Oligoneuron, and the monotypic Oreochrysum, and Semple & al. (1999) formally proposed the expansion of Solidago to reflect this clade. Morphological evidence strongly placed the rayless S. discoidea within Solidago s.s. ...
As currently delimited, Solidagininae are a large (approximately 190 species) subtribe of tribe Astereae. Recent molecular and morphological studies have prompted a new definition of the subtribe, but the lack of absolute morphological synapomorphies raises the possibility that this assemblage may not be monophyletic. Cladistic and likelihood-based analyses were conducted on a nuclear rDNA ITS sequence dataset derived from 23 of the 24 genera included in recent Solidagininae circumscriptions. Cladistic analyses identified two clades entirely composed of proposed Solidagininae genera. The data were not able to support deeper relationships, and these two clades might or might not form one monophyletic lineage. Topology testing indicated compatibility between the taxonomic definition of Solidagininae and molecular data.
... Macrophyllae Semple & J.B. Beck 1 includes one species, Solidago macrophylla Pursh , which was included in subsect. Glomeruliflorae by Semple and Cook (2006), based on RFLP cpDNA results in Zhang (1996) and in the multivariate study of the subsection by Cook et al. (2009). Torrey and Gray (1842) included S. macrophylla Pursh as a synonym under S. squarrosa Muhl. ...
... Solidago than is S. multiradiata. This contradicts the results of the RFLP cp DNA-based study by Zhang (1996; Fig. 1 in Semple et al. 1999 Kiełtyk & Mirek 2014) or are really just alpha level taxonomic proposals that require much more work to document the distinctiveness of the proposed taxon. Based on how taxa of the genus Solidago in North America have apparently rapidly diverged, it seems reasonable to assume that some of the proposed names for European taxa occurring in isolated mountain ranges or distinct habitats might warrant some level of taxonomic recognition within S. virgaurea. ...
A multivariate morphometric study of the goldenrod groups Solidago sect. Solidago and S. subsect. Multiradiatae was conducted to assess the morphological differences among S. virgaurea and the closely related Eurasian species, S. dahurica, S. decurrens (including S. insularis and S. praeflorens), S. horieana, S. kurilensis (including S. nipponica), S. litoralis, S. minutissima, S. pacifica, and S. yokusaiana and the putatively related North American species S. macrophylla and S. leiocarpa, S. multiradiata, and S. spithamaea. Analyses of a matrix of 217 specimens by 34 traits provided support for recognizing 12 species. Statistically, S. macrophylla was not as strongly separated from the Eurasian group of species as was the S. multiradiata group of three species. Some Eurasian taxa were not included in the a priori portions of the analyses. The following new combinations are proposed: Solidago ser. Macrophyllae Semple & J.B. Beck, Solidago sect. Multiradiatae (Semple) Semple & J.B. Beck; Solidago decurrens Loureiro var. insularis (Kitam.) Semple. A neotype is designated for Solidago decurrens and for S. cantonensis.
... Although the systematic relations within Solidago have not been sufficiently resolved, the currently available phylogeny of the genus (Zhang, 1996) allows us to make some assumptions about the evolution of associations between the gall midges and their goldenrod hosts. An emerging pattern from the present study and previous ones (e.g. ...
... It is noteworthy that some of the Asphondylia species use several host plants that are not closely related: for example, A. monacha develops in four Solidago species, each belonging to a different subsection (of Zhang 1996 andSemple &Cook, 2006). Similarly, A. rosulata sp. ...
Reproductive isolation and speciation in herbivorous insects may be accomplished via shifts between host-plant resources: either plant species or plant organs. The intimate association between gall-inducing insects and their host plants makes them particularly useful models in the study of speciation. North American goldenrods (Asteraceae: Solidago and Euthamia) support a rich fauna of gall-inducing insects. Although several of these insects have been the subject of studies focusing on speciation and tritrophic interactions, others remain unstudied and undescribed. Among the latter are at least seven species of the large, cosmopolitan gall midge genus Asphondylia Loew (Diptera: Cecidomyiidae), the taxonomy and biology of which are elucidated here for the first time using morphological, molecular, and life-history data. We describe Asphondylia pseudorosa sp. nov., Asphondylia rosulata sp. nov., and Asphondylia silva sp. nov., and redescribe Asphondylia monacha Osten Sacken, 1869 and Asphondylia solidaginis Beutenmüller, 1907, using morphological characters of adults, immature stages, and galls, as well as sequence data from both nuclear and mitochondrial genes. A neotype is designated for A. solidaginis, the type series of which
is considered lost. We also provide information on the life history of all species, including a description of two
inquilinous cecidomyiids commonly found in the galls, Clinodiplosis comitis sp. nov. and Youngomyia podophyllae (Felt, 1907), and on parasitoid wasps associated with the gall midges. Asphondylia johnsoni Felt, 1908, which was described from an unknown gall on an unknown Solidago host, is assigned to nomina dubia. Our phylogenetic
analyses show that some of the Asphondylia species associated with goldenrods induce two different types of galls during their life cycle, some exhibit host alterations, and some do both. In the absence of reliable morphological differences, recognising species boundaries and deciphering host associations of species must rely heavily on molecular data. Our analysis suggests that radiation in this group has been recent and occurred through shifts among host plants.
... Molecular phylogenies in the tribe initially were based on cpDNA RFLP analyses, mostly of North American (Suh and Simpson 1990;Zanowiac 1991;Morgan and Simpson 1992;Morgan 1993Morgan , 1997Lane et al. 1996;Xiang and Semple 1996;Zhang 1996; see Semple et al. 1999) or Asian genera (Gu et al. 1994;Ito et al. , 1998. Few papers using cpDNA sequence data were published for Astereae Bayer and Cross 2002;Liu et al. 2002;Watanabe et al. 2006;Forest et al. 2007); these data are not cumulative since diff erent markers were used in the studies, which further used a restricted sample of taxa. ...
... This paper summarized his views on the subtribal classifi cation of Astereae for the continent, a modifi cation of Nesom (1994g) now superseded by Nesom and Robinson (2007). Most of the molecular phylogenetic analyses then published were based on RFLP of chloroplast DNA (Suh and Simpson 1990;Morgan and Simpson 1992;Morgan 1993Morgan , 1997Lane et al. 1996;Zhang 1996;Semple et al. 1999), though some of these included ITS data as well (e.g., Morgan 1993Morgan , 1997. Semple et al. (1999Semple et al. ( , 2002 published fl oristic treatments for Ontario goldenrods and asters, respectively, that included phylogenetic discussions. ...
... Molecular phylogenies in the tribe initially were based on cpDNA RFLP analyses, mostly of North American (Suh and Simpson 1990;Zanowiac 1991;Morgan and Simpson 1992;Morgan 1993Morgan , 1997Lane et al. 1996;Xiang and Semple 1996;Zhang 1996; see Semple et al. 1999) or Asian genera (Gu et al. 1994;Ito et al. , 1998. Few papers using cpDNA sequence data were published for Astereae Bayer and Cross 2002;Liu et al. 2002;Watanabe et al. 2006;Forest et al. 2007); these data are not cumulative since diff erent markers were used in the studies, which further used a restricted sample of taxa. ...
... This paper summarized his views on the subtribal classifi cation of Astereae for the continent, a modifi cation of Nesom (1994g) now superseded by Nesom and Robinson (2007). Most of the molecular phylogenetic analyses then published were based on RFLP of chloroplast DNA (Suh and Simpson 1990;Morgan and Simpson 1992;Morgan 1993Morgan , 1997Lane et al. 1996;Zhang 1996;Semple et al. 1999), though some of these included ITS data as well (e.g., Morgan 1993Morgan , 1997. Semple et al. (1999Semple et al. ( , 2002 published fl oristic treatments for Ontario goldenrods and asters, respectively, that included phylogenetic discussions. ...
... Semple et al. (1999) included two distinct species, Solidago glomerata Michx. and Solidago macrophylla Pursh, in the subsection based on cpDNA RFLP analyses (Zhang 1996) and morphology. While some of these taxa are easily recognized, there has been considerable debate in the literature as to the correct taxonomic placement of several of the members of the subsection. ...
... Glomeruliflorae. Zhang (1996) found that S. glomerata and S. macrophylla grouped with S. caesia and S. curtisii in a RFLP cpDNA study of the genus. Not surprisingly, S. glomerata and S. macrophylla were statistiscally strongly supported as distinct in our discriminant analysis of the eight a priori species level groups. ...
A multivariate morphometric study of Solidago subsection Glomeruliflorae (Torr. & A. Gray) Nesom was undertaken on a matrix of 45 characters by 368 plants to assess the morphological differences among the members of the complex and to determine their appropriate taxonomic ranks. The numbers of a priori groups tested were based on taxa recognized in floristic literature, examination of type specimens and 3214 herbarium specimens, and observations of plants in the field and greenhouse. Analyses were performed to determine statistical support for eight a priori species level groups: Solidago albopilosa E.L. Braun, Solidago caesia L., Solidago curtisii Torr. & A. Gray, Solidago flexicaulis L., Solidago glomerata Michx., Solidago lancifolia (Torr. & A. Gray) Chapm., Solidago macrophylla Pursh, and Solidago ouachitensis C.E.S Taylor & R.J. Taylor. Additional analyses were preformed to determine statistical support for varietal level a priori groups within S. caesia and S. curtisii. Statistical support for recognizing all 10 a priori groups varied, although in all analyses. the P values were always well below a of 0.05. Based on the results of the discriminant analyses, the following taxa are recognized: S. albopilosa, S. caesia var. caesia, S. caesia var. zedia, S. curtisii var. curtisii, S. curtisii var. flaccidifolia, S. flexicaulis, S. glomerala, S. lancifolia, S. macrophylla, and S. ouachitensis.
... Alternatively, it is possible that S. gigantea is a younger species, or that it has expanded its distribution into habitats where S. altissima and S. rugosa were already established, thus offering an underused niche to the insects that were associated with these hosts (see Rice, 1987;Berlocher & Feder, 2002). Given the complex and yet unresolved classification of the Solidago species in this group (Zhang, 1996), and the lack of knowledge on their historic distribution in North America (Marks, 1983), these explanations remain speculative. However, it is noteworthy that a shift to S. gigantea has now been documented in four different systems that show varying amounts of gene flow. ...
... These observations imply some involvement of S. rugosa in host shifts and speciation processes in these insects. It appears that despite their presumed distant phylogenetic relations (Zhang, 1996), S. rugosa and the S. altissima species complex (subsection Triplinerviae of Semple & Cook, 2006) are at least physiologically similar with respect to the conditions they offer for the development of gall inducers. ...
The four species of Dasineura Rondani previously reported from goldenrods in North America were revisited, resulting in the synonymy of two species and the redescription of diagnostic characters of adults and galls. Dasineura folliculi Felt is reported from Solidago rugosa Miller and S. gigantea Aiton, and D. carbonaria Felt is reported from Euthamia graminifolia (L.) Nuttall. Both species induce simple leaf-cluster galls on shoot tips of their relative hosts. Dasineura folliculi completes for to five generations, and D. carbonaria completes at least two generations between May and October. Emergence rates were relatively low; they ranged from 36 to 51%, suggesting a large proportion of the larval population does not pupate and instead enters diapause in the ground. D. folliculi galls were attacked by five species of parasitic wasps and by inquilinous and predatory gall midges and caterpillars. Levels of attack by parasitoids increased, whereas levels of attack by inquilines decreased, as the season progressed. Field data and greenhouse experiments indicated that both Dasineura species exhibit monogeny, whereby females produce unisexual broods. Galls resulting from a single oviposition event exclusively yielded single-sex adults, whereas 7–33% of field-collected galls yielded adults of both sexes, suggesting they contained the progeny of more than one female. Gall sex ratios differed between generations of D. folliculi on S. rugosa but not on S. gigantea.
... Hellinsia glenni is certainly behaviourally poorly adapted to galls, as this insect tunnels linearly whereas gall inducers generally restrict their feeding to the gall environment, which requires more circular movement, if any at all. Furthermore, H. glenni is likely physiologically poorly adapted to galls, as these larvae do not normally encounter elevated secondary compounds such as phenolics, which can be three to five times as abundant in E. solidaginis galls as compared to stem tissue [ Abrahamson et al. , 1991 , data for E. solidaginis galls induced on S. altissima , a goldenrod closely related to S. gigantea ( Zhang, 1996 )]. Indeed, in the course of collecting for the current study, H. glenni tunnels were never found in galls, and tunnels that approached the gall turned back into the stem, possibly indicating the gall tissue acts as a feeding deterrent to H. glenni . ...
... Studies of photoassimilate movement and total phenolic levels between galls and stems have shown that the effects of E. solidaginis 's galls are typically very localised. Abrahamson et al. (1991) found that the level of total phenolic compounds in the E. solidaginis galls induced on S. altissima , a goldenrod closely related to S. gigantea ( Zhang, 1996 ), differed significantly from the level of total phenolic compounds 1 cm above or below the gall, and that the total level of phenolic compounds in stems of galled ramets were comparable to those in ungalled ramets. Furthermore, in a study of photoassimilates using C 14 O 2 , McCrea et al. (1985) showed that E. solidaginis galls act as non-mobilising sinks (sieves) rather than as mobilising sinks, which also supports the very localised impact of these galls. ...
Abstract 1. The nutrition hypothesis for the adaptive nature of galls states that gall-inducing insects control the nutrient levels in galls to their own benefit. Although the nutrition hypothesis is widely accepted, there have been few empirical tests of this idea.
2. A novel method is presented for testing the nutrition hypothesis that links manipulation of gall nutrient levels by the gall inducer to herbivore performance. The effects of adaptation and nutritional advantage are separated by using a herbivore that is adapted to a host plant susceptible to galling but one which never enters the gall environment.
3. Hellinsia glenni (Cashatt), a plume moth (Pterophoridae) and one of its host plants provide an excellent system for testing the nutrition hypothesis because H. glenni larvae feed internally on the relatively nutrient-poor stems of a goldenrod, Solidago gigantea, but do not venture into the nutrient-rich galls induced on that plant by a tephritid fly, Eurosta solidaginis. The nutrition hypothesis was tested by transplanting early-instar H. glenni larvae into galls and stems of S. gigantea to determine if the larvae transplanted to galls would perform better compared with those larvae transplanted to stems.
4. The results support the nutrition hypothesis for the adaptive nature of galls. Hellinsia glenni achieved greater final mass in the gall environment compared with the final mass larvae achieved in the stem environment. There was also evidence that the quality of gall tissue is controlled by the gall inducer, which has not been previously demonstrated for mature E. solidaginis galls.
... Alternatively, it is possible that S. gigantea is a younger species, or that it has expanded its distribution into habitats where S. altissima and S. rugosa were already established, thus offering an underused niche to the insects that were associated with these hosts (see Rice, 1987; Berlocher & Feder, 2002). Given the complex and yet unresolved classification of the Solidago species in this group (Zhang, 1996), and the lack of knowledge on their Host-associated speciation in goldenrod gall-midges 735 historic distribution in North America (Marks, 1983), these explanations remain speculative. However, it is noteworthy that a shift to S. gigantea has now been documented in four different systems that show varying amounts of gene flow. ...
... These observations imply some involvement of S. rugosa in host shifts and speciation processes in these insects. It appears that despite their presumed distant phylogenetic relations (Zhang, 1996), S. rugosa and the S. altissima species complex (subsection Triplinerviae of Semple & Cook, 2006 ) are at least physiologically similar with respect to the conditions they offer for the development of gall inducers. ...
Host-associated differentiation (HAD) is considered a step towards ecological speciation and an important mechanism promoting diversification in phytophagous insects. Although the number of documented cases of HAD is increasing, these still represent only a small fraction of species and feeding guilds among phytophagous insects, and most reports are based on a single type of evidence. Here we employ a comprehensive approach to present behavioural, morphological, ecological and genetic evidence for the occurrence of HAD in the gall midge Dasineura folliculi (Diptera: Cecidomyiidae) on two sympatric species of goldenrods (Solidago rugosa and S. gigantea). Controlled experiments revealed assortative mating and strong oviposition fidelity for the natal-host species. Analysis of mitochondrial DNA showed an amount of genetic divergence between the two host-associated populations compatible with cryptic species rather than host races. Lower levels of within-host genetic divergence, gall development and natural-enemy attack in the S. gigantea population suggest this is the derived host.
... presented an intuitive phylogeny of all species of Solidago based on morphological traits and the results of a cpDNA restriction fragment length polymorphisms study (Zhang 1996). The classification scheme of Solidago presented by Nesom (1993) was a critical guide in developing the 2016 intuitive phylogeny, along with modifications presented in the Flora of North America treatment of the genus (Semple and Cook 2006). ...
... Many botanists find Solidago L. taxonomically challenging (Fernald 1950;Croat 1967;Correll and Johnston 1970;Nesom 1993;Voss 1996;Zhang 1996;Cook 2002), a problem stemming from the sheer number of species involved, putative hybridization, and polyploidy. Unfortunately, first-generation DNA sequence data have been of little use in clarifying Solidago species boundaries due to the low observed genetic diversity in the genus. ...
The genus Solidago represents a taxonomically challenging group due to its sheer number of species, putative hybridization, polyploidy, and shallow genetic divergence among species. Here we use a dataset obtained exclusively from herbarium specimens to evaluate the status of Solidago ulmifolia var. palmeri , a morphologically subtle taxon potentially confined to Alabama, Arkansas, Mississippi, and Missouri. A multivariate analysis of both discrete and continuous morphological data revealed no clear distinction between S. ulmifolia var. palmeri and Solidago ulmifolia var. ulmifolia . Solidago ulmifolia var. palmeri ’s status was also assessed with a phylogenomic and SNP clustering analysis of data generated with the “Angiosperms353” probe kit. Neither analysis supported Solidago ulmifolia var. palmeri as a distinct taxon, and we suggest that this name should be discarded. The status of Solidago delicatula (formerly known as Solidago ulmifolia var. microphylla ) was also assessed. Both morphological and phylogenetic analyses supported the species status of S. delicatula and we suggest maintaining this species at its current rank. These results highlight the utility of the Angiosperms353 probe kit, both with herbarium tissue and at lower taxonomic levels. Indeed, this is the first study to utilize this kit to identify genetic groups within a species.
... Many botanists find Solidago L. taxonomically challenging (Fernald 1950;Croat 1967;Correll and Johnston 1970;Voss 1996;Nesom 1993;Zhang 1996;Cook 2002), a problem stemming from the sheer number of species involved, putative hybridization, and polyploidy. ...
The genus Solidago represents a taxonomically challenging group due to its sheer number of species, putative hybridization, polyploidy, and shallow genetic divergence among species. Here we use a dataset obtained exclusively from herbarium specimens to evaluate the status of Solidago ulmifolia var. palmeri, a morphologically subtle taxon potentially confined to Alabama, Arkansas, Mississippi, and Missouri. A multivariate analysis of both discrete and continuous morphological data revealed no clear distinction between S. ulmifolia var. palmeri and Solidago ulmifolia var. ulmifolia. Solidago ulmifolia var. palmeri status was also assessed with a phylogenomic and SNP clustering analysis of data generated with the Angiosperms353 probe kit. Neither analysis supported Solidago ulmifolia var. palmeri as a distinct taxon, and we suggest that this name should be discarded. The status of Solidago delicatula (formerly known as Solidago ulmifolia var. microphylla) was also assessed. Both morphological and phylogenic analyses supported the species status of S. delicatula and we suggest maintaining this species at its current rank. These results highlight the utility of the Angiosperms353 probe kit, both with herbarium tissue and at lower taxonomic levels. Indeed, this is the first study to utilize this kit to identify genetic groups within a species.
... In the present study, the name "goldenrods" refers to the historically congeneric Solidago and Euthamia, which were recently shown to be more distantly related than previously thought (Zhang 1996;Semple & Cook 2006). Six species of goldenrods (Solidago altissima, S. gigantea, S. juncea, S. rugosa, S. nemoralis, and Euthamia graminifolia) were surveyed for galls at least once a week from mid May to mid October 1987 in the following localities in the Cayuga Lake Basin, western NY: Freese Rd. . ...
Goldenrods (Solidago and Euthamia species) are common herbs in the eastern United States that support a large and diverse community of highly specific gall-inducing insects. The majority of these insects are gall midges, of which 16 described species are bud, leaf, stem, rhizome, or flower-head gallers belonging to the large genus Rhopalomyia Rübsaamen. The present work is a taxonomic revision of the goldenrod-associated Rhopalomyia species, which includes a key to the identification of species based on their galls and host plants and descriptions of diagnostic characters for all species. Rhopalomyia lanceolata Felt is designated as a new synonym for R. lobata Felt, and R. albipennis Felt and R. carolina Felt are designated as new synonyms for R. solidaginis Loew. Neotypes are designated for R. hirtipes Osten Sacken and R. solidaginis, and two new species are described—R. gina Dorchin n.sp. and R. guttata Dorchin n.sp. Descriptions include illustrations of galls, male and female morphological characters, and the first description of pupae, where available. New or additional detailed information is also provided on gall structure, phenology, and life history of the gall midges.
... In the present study, the name "goldenrods" refers to the historically congeneric Solidago and Euthamia, which were recently shown to be more distantly related than previously thought (Zhang 1996;Semple & Cook 2006). Six species of goldenrods (Solidago altissima, S. gigantea, S. juncea, S. rugosa, S. nemoralis, and Euthamia graminifolia) were surveyed for galls at least once a week from mid May to mid October 1987 in the following localities in the Cayuga Lake Basin, western NY: Freese Rd. . ...
FIGURES 62 – 69. Rhopalomyia leaf and stem galls. Figs. 62 – 64. Rhopalomyia clarkei galls on Solidago rugosa; 62. Young gall. 63. Mature galls. 64. Galls on underside of leaf. 65. Rhopalomyia clarkei galls on stem and leaf of Solidago altissima (photo by M. Wise). Figs. 66 – 67. Rhopalomyia sp. galls on leaves of Solidago gigantea; 66. Very young galls. 67. Mature gall. Figs. 68 – 69. Rhopalomyia gina galls on leaves of Solidago juncea. 68. Gall on upper side of leaf. 69. Tail-like appendage of gall on underside of leaf.
... In the present study, the name "goldenrods" refers to the historically congeneric Solidago and Euthamia, which were recently shown to be more distantly related than previously thought (Zhang 1996;Semple & Cook 2006). Six species of goldenrods (Solidago altissima, S. gigantea, S. juncea, S. rugosa, S. nemoralis, and Euthamia graminifolia) were surveyed for galls at least once a week from mid May to mid October 1987 in the following localities in the Cayuga Lake Basin, western NY: Freese Rd. . ...
Table of contents Abstract............................................................................................................................................................................... 2 Introduction......................................................................................................................................................................... 2 Materials and methods........................................................................................................................................................ 3 Results and discussion......................................................................................................................................................... 4 Morphology......................................................................................................................................................................... 4 Life history.......................................................................................................................................................................... 5
... Goldenrods (Solidago, Asteraceae) constitute a widespread genus of over 100 species, most of which are native to North America (Zhang 1996). They are insect pollinated and self-incompatible (Melville & Morton 1982) with small, wind-dispersed seeds that do not persist in the soil seed bank (Root 1996). ...
Many long-lived plants are known to prolong dormancy in response to abiotic stresses such as drought. We are unaware, however, of any reports of plants prolonging dormancy in response to biotic stresses such as herbivory. We monitored 140 putative Solidago missouriensis clones (hereafter 'clones') ³13 years before, during and after intense defoliation by the specialist herbivore Trirhabda canadensis. Eight of the clones produced no above-ground growth in the season following defoliation. Though apparently killed, these clones reappeared 1–10 years after they disappeared, with six of them robustly recovering in a single season. We used 38 RAPD markers to test the hypotheses (denoted by H and numbered with subscripts) that territories were recovered by (H1) seedling establishment or (H2) rhizomes. We compared predefoliation and post-recovery genotypes in two clones using the same 38 markers. Our data document the existence of very large clones (60–350 m2 with c. 700–20 000 ramets), and support the hypothesis that recovery is from rhizomes. Within-clone diversity is low, and the pre-defoliation and post-recovery genotypes match. We consider mechanisms that could enable plants entering dormancy with depleted
resources to robustly recover, and the implications of dormancy for avoiding biotic stress such as that induced by T. canadensis.
... Solidago canadensis, S. gigantea, and S. altissima (Asteraceae) are closely related and appear to form a species complex (Melville & Morton, 1982;Zhang, 1996;Semple et al., 1999). Each is common in old fields with S. altissima being the most abundant throughout most of the eastern USA . ...
Host-race formation is promoted by genetic trade-offs in the ability of herbivores to use alternate hosts, including trade-offs due to differential timing of host-plant availability. We examined the role of phenology in limiting host-plant use in the goldenrod gall fly (Eurosta solidaginis) by determining: (1) whether phenology limits alternate host use, leading to a trade-off that could cause divergent selection on Eurosta emergence time and (2) whether Eurosta has the genetic capacity to respond to such selection in the face of existing environmental variation. Experiments demonstrated that oviposition and gall induction on the alternate host, Solidago canadensis, were the highest on young plants, whereas the highest levels of gall induction on the normal host, Solidago gigantea, occurred on intermediate-age plants. These findings indicate a phenological trade-off for host-plant use that sets up the possibility of divergent selection on emergence time. Heritability, estimated by parent-offspring regression, indicated that host-race formation is impeded by the amount of genetic variation, relative to environmental, for emergence time.
Premise of the study:
The phylogenetic relationships among the ca. 138 species of goldenrods (Solidago; Asteraceae) have been difficult to infer due to species richness, and shallow interspecific genetic divergences. This study aims to overcome these obstacles by combining extensive sampling of goldenrod herbarium specimens with the use of a custom Solidago hybrid-sequence capture probe set.
Methods:
A set of tissues from herbarium samples comprising ca. 90% of Solidago species was assembled and DNA was extracted. A custom hybrid-sequence capture probe set was designed, and data from 854 nuclear regions was obtained and analyzed from 209 specimens. Maximum likelihood and coalescent approaches were used to estimate the genus phylogeny for 157 diploid samples.
Key results:
Although DNAs from older specimens were both more fragmented and produced fewer sequencing reads, there was no relationship between specimen age and our ability to obtain sufficient data at the target loci. The Solidago phylogeny was generally well supported, with 88/155 (57%) nodes receiving ≥95% bootstrap support. Solidago was supported as monophyletic, with Chrysoma pauciflosculosa identified as sister. A clade comprising Solidago ericameriodes, Solidago odora, and Solidago chapmanii was identified as the earliest diverging Solidago lineage. The previously segregated genera Brintonia and Oligoneuron were identified as placed well within Solidago. These and other phylogenetic results were used to establish four subgenera and fifteen sections within the genus.
Conclusions:
The combination of expansive herbarium sampling and hybrid-sequence capture data allowed us to quickly and rigorously establish the evolutionary relationships within this difficult, species rich group. This article is protected by copyright. All rights reserved.
Solidago houghtonii Torrey & A. Gray ex Gray is a federally threatened polyploid plant species likely of hybrid origin. Several hypothesized combinations of parental species have been suggested but none have been phylogenetically tested. Additionally, it is unclear whether the species is of single or polytopic origin. To study the evolutionary history of S. houghtonii we sequenced four noncoding cpDNA loci (accD-psaI, psbA-trnH, trnL-trnF, rps16-trnQ), and the ITS and 3′ETS regions for four accessions of S. houghtonii, which span its geographic range, and 25 other species of Solidago including all sympatric species. Polymorphisms within the direct nrDNA sequences of all S. houghtonii accessions indicated the presence of multiple homoeologue types. These were separated by molecular cloning of the 3055 bp 3′ETS — ITS region, allowing us to positively link the ETS and ITS homoeologue types. Phylogenetic analyses of the nuclear and chloroplast datasets revealed incongruent topologies. Analysis of cloned nrDNA sequence data indicated that S. riddellii, S. ptarmicoides, and S. ohioensis have contributed to the nuclear genome of S. houghtonii. Analysis of cpDNA sequence data revealed the presence of multiple insertions/deletions that are shared by all accessions. The unique pattern of cpDNA indels was also recovered in S. gigantea. Phylogenetic analysis of the cpDNA sequence data and coded indels indicate S. gigantea is the maternal genome donor. However, we did not recover a S. gigantea nrDNA sequence type. Taken together, these data reveal both a single origin and a complex pattern of reticulation that is consistent not only with the hypothesized allohexaploid nature of this species, but also with chloroplast capture of cpDNA from an unexpected source through introgression.
For studies of population differentiation and range expansion in the seaside goldenrod, Solidago sempervirens, we identified hypervariable molecular markers by screening genomic libraries enriched for microsatellite motifs. We designed primers that reliably amplified nine polymorphic loci. High polymorphism in a population from Delaware Bay, USA suggests that the loci will be useful in population studies. The success of cross-amplifications in 11 species of Asteraceae varied among loci and did not appear to reflect phylogenetic relationships within Solidago.
Approximately 130 species of goldenrods are native to North America and many occur sympatrically. Such cooccurrence among closely related species raises the question of whether differences among the species in smallscale distribution and growth forms facilitate their co-occurrence. We investigated five goldenrods that frequently co-occur within their native ranges in Pennsylvania USA old fields. We measured goldenrod abundances, soil textures, nutrients, pH, and moisture within 30 old fields, and determined biomass allocation and flower and seed traits for each goldenrod species at a common site. Ordination revealed that Solidago altissima and S. gigantea were associated with fields having circum-neutral soils, whereas Euthamia graminifolia and S. rugosa achieved their highest abundances on acidic soils. Soil clay content and moisture may be associated with a further separation of species as the abundance of S. altissima tended to be higher on well-drained soils while S. gigantea had a tendency to attain its highest abundances on moist soils that had relatively stable moisture levels over time. Euthamia was more likely to be abundant on clay-rich soils while S. rugosa was often associated with soils containing little clay. Solidago juncea tended to associate with droughty soils that underwent marked soil-moisture changes over time. The latter goldenrod had the greatest absolute and relative root mass, the least absolute and relative leaf mass, highest seed-reproductive allocation, and heaviest achenes. In contrast, S. gigantea and Euthamia, which were often associated with more mesic and stable soil moisture conditions, allocated the least to roots and relatively high amounts of mass to leaves. Solidago gigantea, S. altissima, and Euthamia are invasive species across Europe. The species with the highest colonization rate across Europe, S. gigantea, allocated the most to reproduction in our study, while S. altissima, with the second highest colonization rate, was highly clonal producing the most rhizome mass. Life-history trait variation among goldenrods appears to be linked to differences in small-scale distributions and rates of colonization.
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