Subgenus Grammica Research Articles

Palynological studies of some species of the genus Cuscuta with emphasis on taxonomy

The main objective of this research was to conduct a comparative pollen morphological analysis of some representatives of the genus Cuscuta in relation to taxonomy. Investigations of 38 species and 1 variation revealed that not only the number, but the colpi length, as well as the exine ornamentation (at the SEM level), can be regarded as a diagnostic feature. For all the taxa studied, the common zonocolpate type of pollen was observed, represented by three variations: I) colpi long; II) colpi of medium length; III) colpi short. In combination with colpi length, variations in the number of colpi were also identified, such as: I) 3(4)-zonocolpate pollen grains with long colpi (in most of the species studied from subgenera Grammica and Cuscuta, as well as in C. monogyna from the subgenus Monogynella); II) 4(3)-zonocolpate pollen grains, with long colpi (C. cristata from subgenus Grammica); III) 3-zonocolpate pollen grains both with long colpi and colpi of medium length (C. abyssinica from subgenus Cuscuta and C. africana, C. angulata from subgenus Pachystigma); IV) 3-zonocolpate pollen grains with colpi of medium length were noted for species from all four subgenera (C. cassytoides, C. lupuliformis from subgenus Monogynella, C. kotschyana from subgenus Cuscuta, C. nitida from subgenus Pachystigma and C. jalapensis, C. rostrata, C. suaveolens from subgenus Grammica); V) 3–4- or 4-zonocolpate, with short colpi (C. lehmanniana and C. gigantea from subgenus Monogynella respectively); VI) 5–6-zonocolpate, with short colpi (C. reflexa from subgenus Monogynella). Using SEM, three main types of exine ornamentation were also identified: perforate, foveolate and reticulate. The last type in the vast majority of the cases was presented in combination with echinae (rarely with spines or granules). The data revealed that the presence of the significant similarity of palyno-morphological characteristics in representatives of the subgenera Cuscuta, Pachystigma and Grammica, with the exception of the species Lepidanche adpressa (syn. C. compacta), makes it difficult to create an identification key at the level of individual subgenera. Pollen features can be most clearly used only to distinguish the subgenus Monoginella, reflecting its certain isolation within the entire Cuscuta genus. The presence of a highly specialized reticulate exine ornamentation in Lepidanche adpressa (syn. C. compacta), not identified for the pollen of any of the studied taxa of the subgenus Grammica, indicates a significant proximity of this species to the subgenus Monogynella. For the species C. reflexa, the obtained palyno-morphological data are consistent with the previous opinion about the separation of this species, as well as its more advanced position within the genus Cuscuta.

Draining the Swamp Dodders: Systematics of Cuscuta gronovii s.l. (Subgenus Grammica, Sect. Oxycarpae)

Abstract— Cuscuta gronovii species complex (sensu lato, s.l.) is a North American group of species that belong to C. sect. Oxycarpae of subgenus Grammica. Members of C. gronovii s.l. are often associated with riparian or wet habitats, and their delimitation has been problematic historically. DNA sequences from multiple plastid and nuclear ribosomal non-coding regions were used to reconstruct evolutionary relationships among taxa. Molecular phylogeny supported the resurrection of C. saururi, a taxon that has been treated either as C. gronovii var. latiflora or a nomenclatural synonym of C. gronovii var. gronovii. Cuscuta saururi was inferred to be sister to C. cephalanthi, but the relationships among the rest of taxa, although forming a clade with moderate support, remained largely unresolved. Thus, the species limits of C. gronovii s.l. aggregate were also comprehensively investigated through a morphometric study. Plants with 4-merous flowers from New Brunswick and Nova Scotia, Canada, which were previously thought to be C. cephalanthi, were described as a new species, C. acadiana. Cuscuta acadiana, although fertile and producing abundant seeds, is unusual in that the majority of its pollen grains intergrade from zonocolpate, syncolpate, or pantocolpate, with variously shaped or twisted ectoapertures. Cuscuta gronovii var. calyptrata, a widely accepted but virtually unknown taxon, was typified and included in all the analyses. Cuscuta umbrosa was treated as a variety of C. gronovii (var. curta), and C. rostrata was maintained as a species. An identification key, detailed descriptions, illustrations, and a summary of geographical distribution, ecology, and conservation status were provided for all the members of C. gronovii s.l. The only taxon of this species complex found to be weedy, especially in cranberries, was C. gronovii var. gronovii.

Intense proliferation of rDNA sites and heterochromatic bands in two distantly related Cuscuta species (Convolvulaceae) with very large genomes and symmetric karyotypes.

The genome size varies widely among angiosperms but only a few clades present huge variation at a low phylogenetic level. Among diploid species of the genus Cuscuta the genome size increased enormously in at least two independent lineages: in species of subgenus Monogynella and in at least one species (C. indecora) of the subgenus Grammica. Curiously, the independent events lead to similar karyotypes, with 2n = 30 mostly metacentric chromosomes. In this paper we compared the patterns of heterochromatic bands and rDNA sites of C. indecora and C. monogyna, aiming to evaluate the role of these repetitive fractions in these karyotypes. We found out that the large genomes of these species were incremented by a huge number of small heterochromatic CMA+ and DAPI+ bands and 5S and 35 rDNA sites, most of them clearly colocalized with CMA+ bands. Silver nitrate impregnation revealed that the maximum number of nucleoli per nucleus was low in both species, suggesting that some of these sites may be inactive. Noteworthy, the tandem repeats did not generate large bands or sites but rather dozens of small blocks dispersed throughout the chromosomes, apparently contributing to conserve the original karyotype symmetry.

Diversity, evolution and taxonomic significance of fruit in Cuscuta (dodder, Convolvulaceae); the evolutionary advantages of indehiscence

Cuscuta (dodder) is one of the few angiosperm genera characterised by multiple fruit indehiscence/dehiscence modes. Dodder fruit is a capsule with thin pericarp and contains one to four seeds. In some species fruits open at the base (circumscissile dehiscence; DE) while in others they remain closed (indehiscent; IN). IN has evolved multiple times from DE, and is most common in the North American clades of subgenus Grammica. In addition, different forms of “irregular dehiscence” have been anecdotally reported. To determine fruit types and assess homology, we explored fruit development, structure and ultrastructure together with a comparative morphology and character evolution study of other fruit traits. Because it is commonly assumed that IN is associated with a loss of dispersal capability when diaspores contain several seeds, we studied the geographical distribution of species with DE and IN fruits in North America to ascertain whether the latter are less widely distributed than DE species. To determine the putative dispersal advantage(s) of IN we studied the floatability of both capsules and seeds of C. gronovii, a species with IN fruits. Also, in this species we examined the effects of the fruit on seed germination. Pericarp structure/ultrastructure revealed three distinct fruit types: DE, with an abscission zone (AZ); IN without an AZ, and irregularly dehiscent capsules (IrA) which lack an AZ but open because of the thin endocarp cell walls at the fruit base. A fourth functional fruit type, the irregularly dehiscent type B capsules (IrB) was noted, in which dehiscence apparently occurs because of mechanical pressures created among ripening IN fruits within the dense infructescences of the same plant. IrA fruits were posited to be an intermediate evolutionary step between DE and IN fruits; IrA and IrB fruits are not homologous. The only statistically significant fruit trait associated with the dehiscence mode is the size of the interstylar aperture which is larger in IN fruits than in DE capsules. Although not statistically significant, species with IN fruits appear to be more widely distributed in N America that species with DE capsules suggesting an enhanced dispersal capability of IN fruits. Also, species with IN fruits possess a significantly higher northern latitudinal limit in North America. Capsules of C. gronovii were capable of floating for at least one week longer than their seeds indicating that long-distance dispersal by water is possible under certain conditions. Seeds of C. gronovii not removed from their IN fruits exhibited a delayed germination pattern compared to removed seeds, which heightened germination bet hedging and may prevent sibling competition for the same host plants.

Diversity, evolution, and function of stomata bearing structures in Cuscuta (dodders, Convolvulaceae): From extrafloral nectar secretion to transpiration in arid conditions

Abstract Cuscuta includes ca. 200 species of functionally holoparasitic plants grouped in four subgenera: Monogynella, Cuscuta, Pachystigma, and Grammica. Multicellular structures with stomata in Cuscuta are represented by extrafloral nectaries (ENs), reported from the stems of one Monogynella species, and stomatiferous protuberances (SPs), which are non-secretory. These latter structures had been noted on the stems of three Grammica species more than a century ago but entirely forgotten until recently when similar, non-secretory SPs were reported on the flowers of several new Grammica species. Here we study for the first time: (1) the extent of occurrence, diversity and evolution of secretory (ENs) and non-secretory (SPs) multicellular structures in Cuscuta, and (2) the function of SPs. We undertook a character evolution study of ENs and SPs on the stems and flowers of 136 Cuscuta taxa, and examined the structure/ultrastructure of SPs. ENs are inferred as primitive and characterize subg. Monogynella. SPs are derived in the remaining subgenera; they are ubiquitous on the flowers of Cuscuta and Pachystigma, but absent on their stems. Subgenus Grammica species develop two functional types of stems during their life cycle: vegetative, exploratory stems with very low stomatal densities (and no SPs), and reproductive, haustorial stems with numerous SPs. Moreover, 24 species from nine clades of subg. Grammica have evolved morphologically diverse floral SPs with systematic significance. To preliminarily ascertain SP function, we determined in the field the water uptake of Tithonia tubiformis plants parasitized or not by Cuscuta costaricensis, a species with both stem and floral SPs, and the stomatal conductance of dodder stems and flowers, as well as host leaves. Water uptake of parasitized hosts was significantly higher compared to non-parasitized plants, even after host leaves were removed, both during the day and night. The increased water uptake of parasitized hosts and stomatal conductance values suggest a transpiration role for the SPs, which is also confirmed by their lacunar structure. Grammica species with floral SPs grow in arid areas or characterized by a pronounced dry season during flowering/fruiting, which suggests that SPs may have evolved to stimulate the host water uptake during these phenophases.

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

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. 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. 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. 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.

More Problems Despite Bigger Flowers: Systematics of <I>Cuscuta tinctoria</I> Clade (subgenus <I>Grammica</I>, Convolvulaceae) with Description of Six New Species

Abstract Cuscuta tinctoria clade, the second largest infrageneric group of subgenus Grammica, includes 14 species that are centered in Mexico and adjacent regions, but also two species that are found in Australia, presumably as a result of long-distance dispersal. Flowers, pollen, and stigmas are among the largest in Cuscuta, with plants being xenogamous or facultatively xenogamous. In this clade, the convergent evolution of morphological traits, especially those associated with the calyx and gynoecium/ capsule, has obscured the identity of some species and has hindered previous efforts to determine their limits. Basic morphology, scanning electron microscopy and sequence data from the nuclear internal transcribed spacer (ITS) and the plastid trnL-F region were used to reconstruct the phylogeny, gain a better understanding of the evolutionary history, and determine species boundaries. Overall, species were grouped in five subclades, Based on their morphological and molecular similarity, C. tinctoria, C. a...

Plastid genome evolution across the genus Cuscuta (Convolvulaceae): two clades within subgenus Grammica exhibit extensive gene loss.

The genus Cuscuta (Convolvulaceae, the morning glory family) is one of the most intensely studied lineages of parasitic plants. Whole plastome sequencing of four Cuscuta species has demonstrated changes to both plastid gene content and structure. The presence of photosynthetic genes under purifying selection indicates that Cuscuta is cryptically photosynthetic. However, the tempo and mode of plastid genome evolution across the diversity of this group (~200 species) remain largely unknown. A comparative investigation of plastid genome content, grounded within a phylogenetic framework, was conducted using a slot-blot Southern hybridization approach. Cuscuta was extensively sampled (~56% of species), including groups previously suggested to possess more altered plastomes compared with other members of this genus. A total of 56 probes derived from all categories of protein-coding genes, typically found within the plastomes of flowering plants, were used. The results indicate that two clades within subgenus Grammica (clades ‘O’ and ‘K’) exhibit substantially more plastid gene loss relative to other members of Cuscuta. All surveyed members of the ‘O’ clade show extensive losses of plastid genes from every category of genes typically found in the plastome, including otherwise highly conserved small and large ribosomal subunits. The extent of plastid gene losses within this clade is similar in magnitude to that observed previously in some non-asterid holoparasites, in which the very presence of a plastome has been questioned. The ‘K’ clade also exhibits considerable loss of plastid genes. Unlike in the ‘O’ clade, in which all species seem to be affected, the losses in clade ‘K’ progress phylogenetically, following a pattern consistent with the Evolutionary Transition Series hypothesis. This clade presents an ideal opportunity to study the reduction of the plastome of parasites ‘in action’. The widespread plastid gene loss in these two clades is hypothesized to be a consequence of the complete loss of photosynthesis. Additionally, taxa that would be the best candidates for entire plastome sequencing are identified in order to investigate further the loss of photosynthesis and reduction of the plastome within Cuscuta.

To attract or to repel? Diversity, evolution and role of the “most peculiar organ” in the Cuscuta flower (dodder, Convolvulaceae)—the infrastaminal scales

Cuscuta (dodders) is the only genus in Convolvulaceae and the Solanales more broadly in which corolla-stamen structures called infrastaminal scales (IFS) have reached a high degree of elaboration and diversification in a great number of species. Historically, morphological diversity of IFS has provided some of the most useful taxonomic characters at the species-level. However, their function has not been determined. We have performed a comparative study of the IFS in 147 Cuscuta taxa using light, scanning and transmission electron microscopy, and results were analyzed in relation to a Cuscuta phylogeny obtained from a combined analysis of nuclear ITS and plastid trnL-F sequences. The morphology and histochemistry of scales and/or trichomes on the staminal filaments were also examined in several other Convolvulaceae genera to provide a preliminary foundation for homology interpretation. To test the hypothesis that the role of IFS in Cuscuta is connected to sexual reproduction, we analyzed the correlations between scale characters and previously published values for pollen/ovule ratios and the number of stomata found in the nectary at the base of the ovary. While the correlations at the level of the entire genus were very low, subgeneric partitions revealed a strong relationship between scale characters and sexual reproduction indicators in subgenus Monogynella. However, this connection declined drastically in the subgenera Cuscuta and Grammica. Our results strongly suggest that scales in Cuscuta evolved in connection to a modification of their function in the flower: from nectar protection and holding in the first diverged subgenus Monogynella, to ovary/ovule protection against herbivorous insects in the derived subgenera Cuscuta and Grammica. Thus at least in the case of the latter subgenera, the protective/repellent role of the IFS may conflict with the attractant/rewarding function of the nectary found at the base of the ovary. In subgenus Monogynella, IFS fimbriae are similar to uniseriate glandular hairs, with the secretory cells entirely exposed. In the subgenera Grammica and Cuscuta, the fimbriae become more complex, with an internal distal laticifer, and a precise mechanism of latex release. Our study elaborates further on the development and ultrastructure of scales in C. gronovii, and provides details on the anatomy of the IFS in other species. The new morphology and micromorphology data confirm the significance of these structures for species-level systematics.

Diversity and evolution of pollen-ovule production in Cuscuta (dodders, Convolvulaceae) in relation to floral morphology

Cuscuta (dodder, Convolvulaceae) is a genus of about 200 species of obligate stem parasites with subcosmopolitan distribution. The diversity of pollen and ovule production was surveyed in 128 species and ten varieties. Taxa were assigned to Cruden’s mating system categories based on their pollen-ovule ratios. Variation and correlations among floral characters were analyzed using regression and ANOVA, while the mating system categories were subjected to a linear discriminant and canonical variates analysis to assess their cohesiveness. Our data strongly suggest that most Cuscuta species possess a wide range of mixed-mating systems. Whereas four ovules develop in each flower, pollen production varies over three orders of magnitude. Several Cuscuta taxa are highly outcrossing, but no species could be identified that are exclusively selfing. The transition from the one-style flowers of subg. Monogynella to the two-style flowers of subgenera Cuscuta and Grammica, and from simultaneous to sequential maturation of the two stigmas in the latter subgenus, has decreased the role of herkogamy as a facilitator of outcrossing. These evolutionary changes are associated with an increase of species richness in subgenus Cuscuta, and especially in subgenus Grammica. Morphological features were not individually found to have a strong correlation to the mating system, but in general, larger corollas and stigmas were associated with greater pollen-ovule ratios. Cuscuta presents some puzzling results when considered in light of the sex allocation theory, as only some infrageneric lineages demonstrate the predicted pollen size-number tradeoff, while Cuscuta gracillima complex (in subgenus Grammica) displays an unexpected negative relationship between pollen size and style length. The relationship between host range and mating system is discussed, prompting further research into the co-evolution of pollination systems and life history traits between parasites and their host species.

Systematics of “horned” dodders: phylogenetic relationships, taxonomy, and two new species within the Cuscuta chapalana complex (Convolvulaceae)

The Cuscuta chapalana complex (Convolvulaceae) is a recently circumscribed clade within the subgenus Grammica that includes several little-known species from Mexico and Central and northern South America characterized by the presence of peculiar subapical multicelullar appendages on the corolla and often on the calyx lobes. Basic morphology, scanning electron microscopy, and DNA sequence data from the plastid trnL–F region and the 26S rDNA and internal transcribed spacer nuclear regions were used to investigate the phylogenetic relationships within the group and to test the species limits. Multicellular appendages are horn-like or tubular in shape and bear one or more stomata at their tips; hence the vernacular name “horned” dodders. Eight lineages were reconstructed; two of them, both from Mexico, were described as new species: Cuscuta bonafortunae Costea & I. García, sp. nov., and Cuscuta carnosa Costea & Stefanović, sp. nov. Cuscuta pringlei is redefined as a variety of Cuscuta strobilacea . A taxonomic treatment with an identification key, descriptions, and illustrations is provided.

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.

Reticulate evolution in the parasitic genusCuscuta(Convolvulaceae): over and over againThis paper is one of a selection of papers published in the Special Issue on Systematics Research.

The frequency and relative importance of hybridization in plants has been an area of intense debate. Although this evolutionary phenomenon has received considerable attention from plant biologists, there are no well-supported cases of reticulate evolution involving parasitic plants, to date. Recent molecular phylogenetic analyses revealed that the subgenus Grammica , the largest and most diverse group of the stem-parasitic genus Cuscuta (dodder), consists of 15 major clades. We describe here five cases of strongly supported discordance between phylogenies derived from plastid and nuclear data, and interpret them as results of five independent hybridization events. Three of these cases could represent relatively recent reticulations, as each of them involves more closely related species, always confined within the same major clade as their putative parental species, and are currently sympatric or parapatric with them. The two remaining cases involve species whose potential progenitors are derived from different major groups of Grammica, and which are allopatric in their present distribution. A series of statistical tests was conducted to assess and further explore the significance of this phylogenetic incongruence. Alternative explanations for discordant gene topologies are explored. Cuscuta liliputana sp. nov., a new Mexican species of hybrid origin is described.

Systematics and plastid genome evolution of the cryptically photosynthetic parasitic plant genus Cuscuta(Convolvulaceae)

BackgroundThe genus Cuscuta L. (Convolvulaceae), commonly known as dodders, are epiphytic vines that invade the stems of their host with haustorial feeding structures at the points of contact. Although they lack expanded leaves, some species are noticeably chlorophyllous, especially as seedlings and in maturing fruits. Some species are reported as crop pests of worldwide distribution, whereas others are extremely rare and have local distributions and apparent niche specificity. A strong phylogenetic framework for this large genus is essential to understand the interesting ecological, morphological and molecular phenomena that occur within these parasites in an evolutionary context.ResultsHere we present a well-supported phylogeny of Cuscuta using sequences of the nuclear ribosomal internal transcribed spacer and plastid rps2, rbcL and matK from representatives across most of the taxonomic diversity of the genus. We use the phylogeny to interpret morphological and plastid genome evolution within the genus. At least three currently recognized taxonomic sections are not monophyletic and subgenus Cuscuta is unequivocally paraphyletic. Plastid genes are extremely variable with regards to evolutionary constraint, with rbcL exhibiting even higher levels of purifying selection in Cuscuta than photosynthetic relatives. Nuclear genome size is highly variable within Cuscuta, particularly within subgenus Grammica, and in some cases may indicate the existence of cryptic species in this large clade of morphologically similar species.ConclusionSome morphological characters traditionally used to define major taxonomic splits within Cuscuta are homoplastic and are of limited use in defining true evolutionary groups. Chloroplast genome evolution seems to have evolved in a punctuated fashion, with episodes of loss involving suites of genes or tRNAs followed by stabilization of gene content in major clades. Nearly all species of Cuscuta retain some photosynthetic ability, most likely for nutrient apportionment to their seeds, while complete loss of photosynthesis and possible loss of the entire chloroplast genome is limited to a single small clade of outcrossing species found primarily in western South America.

Delimitation of major lineages withinCuscutasubgenusGrammica(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.

Down the Slippery Slope: Plastid Genome Evolution in Convolvulaceae

Cuscuta (dodder) is the only parasitic genus found in Convolvulaceae (morning-glory family). We used long PCR approach to obtain large portions of plastid genome sequence from Cuscuta sandwichiana in order to determine the size, structure, gene content, and synteny in the plastid genome of this Cuscuta species belonging to the poorly investigated holoparasitic subgenus Grammica. These new sequences are compared with the tobacco chloroplast genome, and, where data are available, with corresponding regions from taxa in the other Cuscuta subgenera. When all known plastid genome structural rearrangements in parasitic and nonparasitic Convolvulaceae are considered in a molecular phylogenetic framework, three categories of rearrangements in Cuscuta are revealed: plesiomorphic, autapomorphic, and synapomorphic. Many of the changes in Cuscuta, previously attributed to its parasitic mode of life, are better explained either as plesiomorphic conditions within the family, i.e., conditions shared with the rest of the Convolvulaceae, or, in most cases, autapomorphies of particular Cuscuta taxa, not shared with the rest of the species in the genus. The synapomorphic rearrangements are most likely to correlate with the parasitic lifestyle, because they represent changes found in Cuscuta exclusively. However, it appears that most of the affected regions, belonging to all of these three categories, have probably no function (e.g., introns) or are of unknown function (a number of open reading frames, the function of which, if any, has yet to be discovered).

Pollen morphology of Cuscuta (Convolvulaceae) in Taiwan.

The pollen morphology of five taxa of Cuscuta, C. australis, C. campestris, C. chinensis, C. japonica var. formosana and C. japonica var. japonica (Convolvulaceae) in Taiwan was investigated by using LM, SEM and TEM. The pollen of Cuscuta is three to six colpate, zonocolpate or pantocolpate colpus with granules, tectum with scabrate processes, ektexine reticulate or finely reticulate, and endexine thinner than ektexine. Two distinct pollen types of Cuscuta spp. in Taiwan are recognized. Type 1 is small, and has colpus with granules, and ektexine finely reticulate, including C. australis, C. campestris and C. chinensis. Type 2 is medium in size, and has colpus with granules, scabrate processes on surface of granule and ektexine reticulate, including C. japonica var. formosana and C. japonica var. japonica. Pollen types correspond with Yuncker's subdivision of the genus Cuscuta, type 1 for subgenus Grammica and type 2 for subgenus Monogyna. Meanwhile, pollen characteristics are also helpful in identifying species.

Chromosome divergence in the genusCuscutaand its systematic implications

SUMMARYCuscuta L. plants are holoparasites with high chromosome divergence. Species representing the three subgenera of the genus were checked for their chromosome number, length and behaviour. Chromosome numbers range between 2n = 8 and 2n = 60, and their size, although roughly uniform within species, varies between 1–23μm. On the whole, size or number of Cuscuta chromosomes cannot be used as diagnostic criteria, except for very few species with unique numbers or sizes. Some associations between certain systematic entities and chromosome characters can be inferred. Species of the Old World of subgenus Cuscuta Engelm. are characterized by inverted meiosis and apparently by holocentric chromosomes, whereas species of the two other subgenera, subgenus Motiogyna Engelm. and subgenus Grammica Yuncker, undergo the regular process of meiosis and have monocentric chromosomes. The presence of holocentric chromosomes does not seem to be associated with their number.

Unusual chromosome separation in meiosis of Cuscuta L.

This study suggests that precocious division of centromeres and probably inverted meiosis (i.e., the rare occasion in meiosis when the equatorial division precedes the reductional division) occur not only in Cuscuta babylonica Choisy but also in several other dodders, regardless of their chromosome number. It was found in four additional species, all of which belong to subgenus Cuscuta. At least two other species (of subgenus Grammica and subgenus Monogyna) undergo the usual process. Therefore, Cuscuta warrants closer study and may provide a better understanding of inverted meiosis as well as the regular process of meiosis.Key words: inverted meiosis, precocious division of centromere, Cuscuta spp.