Mihai Costea

Delimitation of major lineages within Cuscuta subgenus Grammica (Convolvulaceae) using plastid and nuclear DNA sequences

Subgenus Grammica, the largest and most diverse group in the parasitic genus Cuscuta, includes 130 species distributed primarily throughout the New World, with Mexico as its center of diversity. To circumscribe the subgenus and assess the relationships among its major lineages, we conducted the first phylogenetic study of Grammica using plastid trnL-F and nrITS sequences from a wide taxonomic sampling covering its morphological, physiological, and geographical diversity. With the exception of one species belonging elsewhere, the subgenus was found to be monophyletic. The results further indicate the presence of 15 well-supported major clades within Grammica. Some of those lineages correspond partially to earlier taxonomic treatments, but the majority of groups are identified in this study for the first time. The backbone relationships among major clades, however, remain weakly supported or unresolved in some cases. The phylogenetic results indicate that the fruit dehiscence character is homoplastic, thus compromising its value as a major taxonomic and evolutionary feature. While several striking cases of long-distance dispersal are inferred, vicariance emerges as the most dominant biogeographical pattern for Cuscuta. Species placed within one of the clades with a predominantly South American distribution are hypothesized to have substantially altered plastid genomes.

The Biology of Invasive Alien Plants in Canada. 3. Amaranthus tuberculatus (Moq.) Sauer var. rudis (Sauer) Costea & Tardif

This annual dioecious weed was found in 2002 and 2003 infesting soybean fields in southwestern Ontario, and it was collected in 1992 from waste places in British Columbia. It is a major weed problem in field crops in the mid-western United States, where it has become increasingly difficult to control during the past 10 yr. Morphological differences between Amaranthus tuberculatus var. rudis and var. tuberculatus are presented. A review of the biological information published is provided. Plants exhibit high phenotypic plasticity and genetic variability. Emergence is prolonged, growth rapid, and female plants produce a large number of viable seeds that contribute to a persistent seed bank. Amaranthus tuberculatus var. rudis has developed multiple resistance to triazine and acetolactate synthase- and protoporphyrinogen-inhibiting herbicides. Airborne pollen can travel significant distances and A. tuberculatus var. rudis may hybridize with other noxious Amaranthus spp. transferring herbicide resistance or ot...

The biology of Canadian weeds. 133. Cuscuta campestris yuncker, C. gronovii willd. ex Schult., C. umbrosa beyr. ex hook., C. epithymum (L.) L. and C. epilinum Weihe

Cuscuta spp. (dodders) are rootless, holoparasitic herbs with filiform stems attached to the host by numerous haustoria. In Canada, Cuscuta gronovii is the most common native species of the genus followed by Cuscuta campestris and C. umbrosa. Cuscuta epithymum and C. epilinum, both introduced species in Canada, occur occasionally. Infestation by Cuscuta spp. can result in serious yield losses and dodders are listed as noxious weeds in British Columbia, Ontario and Quebec, and as restricted weeds in Alberta. These plants have evolved special adaptations to ensure their success: germination occurs late in the season when potential hosts are already established; seedlings selectively forage in plant communities and they may survive relatively long periods during the autotrophic stage. Invasion occurs via extremely elaborate mechanisms designed to match the biological processes of their host and bypass defense mechanisms. The principal means of dispersal of Cuscuta weeds world-wide (including Canada) has been...

Cuscuta jepsonii (Convolvulaceae): an invasive weed or an extinct endemic?

Despite their ecological significance, parasitic plants face more conservation challenges than do autotrophic plants. This is especially true for the groups that include weedy or invasive species such as Cuscuta. While approximately half of the Cuscuta (dodders) species may require conservation measures, the genus as a whole is sometimes posted on governmental lists of noxious or quarantine weeds. Our study challenges this stereotype and uses the case of C. jepsonii (Jepsons dodder) to illustrate the precarious biodiversity and conservation status faced by many dodder species. Until now, Jepsons dodder has been known only from its type collection. Consequently, its phylogenetic affinities, morphological variation, and ecology have remained unknown, and the species is currently ambiguously considered either synonymous to the invasive North American weed C. indecora or to an extinct endemic from California. Using molecular data from newly found collections, we infer that C. jepsonii belongs to C. californica species complex, instead of C. indecora clade. Also, we discuss the conservation of this species within the broader biological and ecological context of Cuscuta in general.

The biology of Canadian weeds. 131. Polygonum aviculare L.

A review and assessment of biological information as well as personal data are provided for Polygonum aviculare in Canada. The species has been revised taxonomically and the six subspecies that occur in Canada are presented. Three of the subspecies, P. aviculare subsp. aviculare, P. aviculare subsp. neglectum and P. aviculare subsp. depressum are weeds introduced to Canada from Europe. A fourth subspecies, P. aviculare subsp. buxiforme is apparently native to North America. The geographical distribution of the latter four subspecies is very wide. Plants exhibit a high phenotypic plasticity and genetic variability, and they easily adapt to a multitude of agrestal and ruderal habitats. The seeds have a variable dormancy and polymorphic germination as a result of heterocarpy, genetic and environmental factors. In other areas (Europe), the species has developed resistance to triazines. Plants are hosts to nematodes, viruses, and fungi that also affect cultivated plants. Key words: POLAV, ecology, distribution...

Phylogeny, character evolution, and biogeography of Cuscuta (dodders; Convolvulaceae) inferred from coding plastid and nuclear sequences

PREMISE OF THE STUDY The parasitic genus Cuscuta, containing some 200 species circumscribed traditionally in three subgenera, is nearly cosmopolitan, occurring in a wide range of habitats and hosts. Previous molecular studies, on subgenera Grammica and Cuscuta, delimited major clades within these groups. However, the sequences used were unalignable among subgenera, preventing the phylogenetic comparison across the genus. METHODS We conducted a broad phylogenetic study using rbcL and nrLSU sequences covering the morphological, physiological, and geographical diversity of Cuscuta. We used parsimony methods to reconstruct ancestral states for taxonomically important characters. Biogeographical inferences were obtained using statistical and Bayesian approaches. KEY RESULTS Four well-supported major clades are resolved. Two of them correspond to subgenera Monogynella and Grammica. Subgenus Cuscuta is paraphyletic, with section Pachystigma sister to subgenus Grammica. Previously described cases of strongly supported discordance between plastid and nuclear phylogenies, interpreted as reticulation events, are confirmed here and three new cases are detected. Dehiscent fruits and globose stigmas are inferred as ancestral character states, whereas the ancestral style number is ambiguous. Biogeographical reconstructions suggest an Old World origin for the genus and subsequent spread to the Americas as a consequence of one long-distance dispersal. CONCLUSIONS Hybridization may play an important yet underestimated role in the evolution of Cuscuta. Our results disagree with scenarios of evolution (polarity) previously proposed for several taxonomically important morphological characters, and with their usage and significance. While several cases of long-distance dispersal are inferred, vicariance or dispersal to adjacent areas emerges as the dominant biogeographical pattern.

Systematics of Cuscuta chinensis species complex (subgenus Grammica, Convolvulaceae): evidence for long-distance dispersal and one new species

Cuscuta chinensis species complex is a small clade from subgenus Grammica (Cuscuta, Convolvulaceae). Many species of this clade exhibit crest- or dome-like multicellular appendages with stomata on the midvein/carena of calyx lobes. Basic morphology, scanning electron microscopy, and DNA sequence data from the plastid trnL-F region and the nuclear internal transcribed spacer (ITS) were used to investigate the phylogenetic relationships within the group and test the species limits. Based on their morphological and molecular similarly, C. chinensis and C. applanata represent one single species, and the latter was retained as a variety of the former. While the clade is centered in the southern USA and Mexico, C. chinensis var. chinensis has a disjunct distribution in Australia and Asia, which is likely the result of relatively recent long-distance dispersal. Cuscuta alata from Mexico, previously considered synonymous to C. applanata, was found to be a distinct species based on both molecular and morphological evidence. Cuscuta potosina var. potosina and var. globifera segregate into two different subclades of the C. chinensis species complex. Because these two taxa are also different morphologically, var. globifera is described as a new species, C. azteca. Cuscuta campestris, an almost cosmopolitan weed species from another clade (“Clade B”; “C. pentagona complex”), is commonly misidentified in Asia as C. chinensis, which raises questions about the identity of the Cuscuta plants used in widely commercialized medicinal herbal mixtures. A taxonomic treatment with an identification key, descriptions, and illustrations is provided.

Diversity and evolution of the gynoecium in Cuscuta (dodders, Convolvulaceae) in relation to their reproductive biology: two styles are better than one

The gynoecium of 122 species and 14 varieties of Cuscuta (dodders) was investigated by light and scanning electron microscopy to assess its diversity and evolution and to provide a morphological foundation for understanding the different reproductive strategies encountered in the genus. Data were optimized into a consensus tree constructed from three large-scale molecular phylogenies of the genus based on nuclear ITS and plastid trnL-F sequences. The number of styles combined with the stigma shape are the only floral/fruit characters that enable the separation of Cuscuta subgenera. In addition, gynoecium morphology is useful for delimiting species in some clades. The one-style gynoecium of subg. Monogynella is mostly likely ancestral whereas gynoecia with two styles are derived in subgenera Cuscuta and Grammica. Gynoecia with two styles encountered in the latter subgenera provide a greater morphological complexity and flexibility for various reproductive strategies. In subg. Cuscuta, both the equal styles and stigmas continue to elongate and modify their position after the flowers open, until pollination occurs. In subg. Grammica, the two unequal styles may cause a spatial separation of the sexes in the flower, herkogamy, while the two stigmas mature sequentially and have a differential timing of their receptivity for pollen. A nectary consisting of a ring of modified stomata at the base of the ovary, the equivalent of the hypogynous nectary disc present in many Convolvulaceae, was observed for the first time in all Cuscuta species. The vasculature of the styles is reduced, represented mostly by phloem; xylem is present only in subg. Monogynella. Some gynoecial characters, for example papillae diameter, stigma surface area, stigma width, and style width were moderately correlated with pollen volume, pollen polar–equatorial ratio and tectum perforation. Gynoecium features suggest that Cuscuta is allied with the “bifid clade” (Dicranostyloideae) in Convolvulaceae.

Untangling the Systematics of Salt Marsh Dodders: Cuscuta pacifica, a New Segregate Species from Cuscuta salina (Convolvulaceae)

Abstract The salt marsh dodders, Cuscuta salina, have been historically delimited as a morphologically variable assemblage of inbreeding forms that parasitize hosts growing in alkaline or saline habitats from western North America. This morphological diversity has been traditionally classified into three varieties: salina, major, and papillata. A morphometric analysis of floral characters and a molecular study using both plastid and nuclear DNA sequences strongly support the segregation of a new species, Cuscuta pacifica Costea and M. A. R. Wright, from C. salina. The new species corresponds to a lineage that includes varieties major and papillata, whereas C. salina is limited essentially to its type variety. Cuscuta pacifica and C. salina are sister species that have only a small area of parapatry in lower California, where they are ecologically and reproductively separated. Cuscuta salina occurs mostly in inland vernal pools and salt flats of Arizona, California, Nevada, Utah, Baja California, and Sonora, and grows primarily on Frankenia and Suaeda. Cuscuta pacifica can be found in salt marshes from the southcentral Pacific coast of California north into British Columbia, parasitic especially on Salicornia and Jaumea carnosa. Cuscuta salina var. papillata (Yunck.) Costea and M. A. R. Wright, parasitic on hosts that grow in coastal interdunes, falls within the range of variation of C. pacifica, where it is transferred.

A Phylogenetically Based Infrageneric Classification of the Parasitic Plant Genus Cuscuta (Dodders, Convolvulaceae)

Abstract— Cuscuta (dodders, Convolvulaceae) is one of the largest and most economically important lineages of parasitic plants. The genus has a sub-cosmopolitan distribution with more than 75% of the species diversifying in the New World. The last monograph, published by Truman George Yuncker in 1932, provided a solid species-level taxonomic foundation. However, as revealed by recent phylogenetic studies, its infrageneric classification has been in great need of a taxonomic reappraisal, mainly because the morphological characters used in the previous classifications have been greatly affected by convergent evolution. Several recent phylogenetic and character evolution studies with broad sampling, as well as species-level revisions, have illustrated the deficiencies of previous classifications and provided an explicit and robust phylogenetic framework. Here we propose a new phylogenetic classification that places all 194 currently accepted species of Cuscuta into four subgenera and 18 sections. Sections have a strong morphological and biogeographical predictive value and include from one to 31 species. Thirteen section names are new or applied for the first time at the sectional rank: Babylonicae (Yunck.) M. A. García, Subulatae (Engelm.) Costea & Stefanović, Obtusilobae (Engelm.) Costea & Stefanović, Prismaticae (Yunck.) Costea & Stefanović, Ceratophorae (Yunck.) Costea & Stefanović, Umbellatae (Yunck.) Costea & Stefanović, Gracillimae Costea & Stefanović, Californicae (Yunck.) Costea & Stefanović, Indecorae (Yunck.) Costea & Stefanović, Oxycarpae (Engelm. ex Yunck.) Costea & Stefanović, Racemosae (Yunck.) Costea & Stefanović, Partitae Costea & Stefanović, and Denticulatae (Yunck.) Costea & Stefanović. An identification key to sections is included together with an overview of morphology, geographical distribution, taxonomic notes, and lists of included species.