Plastome variation and phylogeny of Taxillus (Loranthaceae).

Several molecular phylogenetic studies of the mistletoe family Loranthaceae have been published such that now the general pattern of relationships among the genera and their biogeographic histories are understood. Less is known about species relationships in the larger (> 10 species) genera. This study examines the taxonomically difficult genus Taxillus composed of 35-40 Asian species. The goal was to explore the genetic diversity present in Taxillus plastomes, locate genetically variable hotspots, and test these for their utility as potential DNA barcodes. Using genome skimming, complete plastomes, as well as nuclear and mitochondrial rDNA sequences, were newly generated for eight species. The plastome sequences were used in conjunction with seven publicly available Taxillus sequences and three sequences of Scurrula, a close generic relative. The Taxillus plastomes ranged from 121 to 123 kbp and encoded 90-93 plastid genes. In addition to all of the NADH dehydrogenase complex genes, four ribosomal genes, infA and four intron-containing tRNA genes were lost or pseudogenized in all of the Taxillus and Scurrula plastomes. The topologies of the plastome, mitochondrial rDNA and nuclear rDNA trees were generally congruent, though with discordance at the position of T. chinensis. Several variable regions in the plastomes were identified that have sufficient numbers of parsimony informative sites as to recover the major clades seen in the complete plastome tree. Instead of generating complete plastome sequences, our study showed that accD alone or the concatenation of accD and rbcL can be used in future studies to facilitate identification of Taxillus samples and to generate a molecular phylogeny with robust sampling within the genus.

Floral anatomy of the plant Psittacanthus schiedeanus (Loranthaceae) / Anatomía floral de la planta Psittacanthus schiedeanus (Loranthaceae)

Loranthaceae hemiparasitic family comprises 76 genera and about 1 050 species distributed in temperate and tropical regions.The subtribe Psittacanthinae contains 14 genera of neotropical mistletoe including Psittacanthus with over 120 species, characterized by large, brightly colored (red, orange, yellow) flowers that are mostly pollinated by hummingbirds. During the 20th century, a number of morphological and embryological studies were conducted mainly on Old World Loranthaceae genera. More recently, attention has been focused on neotropical Psittacanthinae where among the 14 genera, floral anatomy has been examined in only seven.The aim of this study is to describe the floral anatomy of Psittacanthus schiedeanus and compares the results with those derived from related mistletoe, interpreting the variation of the floral characters of the calyculus, nectary, gynoecium and from floral dissections and serial histological sections, detailing the structure of androecium and gynoecium and anthers in the context of the new phylogenetic information. Flowers of P. schiedeanus at different developmental stages were examined using stained serial sections visualized with light microscopy. These flowers have a vascularized, cupular pedicel fused to a bracteole, a non-vascularized calyculus, an annular nectary, a unilocular gynoecium with a single central mamelon and an androecium formed by epipetalous septate stamens. The morphological comparison of pedicel, bracteole and calyculus provides support for the interpretation of the calyculus as a reduced calyx. The annular nectary seems to be a character shared by the entire subtribe Psittacanthinae, which distinguishes it from Ligarinae which has stylar nectary. The unilocular gynoecium formed by a single central structure is a character shared with other genera in Psittacanthinae except Tripodanthus. The androecium is composed of dithecal, tetrasporangiate stamens with septate locules that are here considered an adaptation for pollen releasing over an extended time period rather than previous suggestions that they result from evolutionary pressure to reduce anther size or to facilitate the nutrition of microspores in large anthers.

La familia hemiparásita Loranthaceae comprende 76 géneros y aproximadamente 1 050 especies distribuidas en regiones templadas y tropicales. La subtribu Psittacanthinae contiene 14 géneros de muérdagos neotropicales que incluyen Psittacanthus con más de 120 especies, caracterizadas por presentar flores grandes de colores brillantes (rojo, naranja y amarillo) que son polinizadas principalmente por colibríes. Durante el siglo XX se desarrollaron una serie de estudios morfológicos y embriológicos de géneros de Loranthaceae del Viejo Mundo. Recientemente, la atención se ha centrado en la subfamilia neotropical Psittacanthinae, en donde de los 14 géneros que la conforman, la anatomía floral se ha examinado solamente en siete. El objetivo de este estudio es describir la anatomía floral de Psittacanthus schiedeanus y comparar los resultados con los de otros muérdagos relacionados, interpretando la variación de los caracteres florales del calículo, nectario, gineceo y anteras en el contexto de la nueva información filogenética. Flores de P. schiedeanus en diferentes estados de desarrollo fueron examinadas mediante secciones seriadas teñidas utilizando microscopía óptica. Estas flores tienen un pedículo vascularizado y cupular fusionado con una bracteola, un cáliz no vascularizado, un nectario anular, un gineceo unilocular con un solo mamelón central y un androceo formado por estambres septados epipétalos. La comparación morfológica de pedicelo, bracteola y calículo proporciona apoyo para la interpretación del calículo como un cáliz reducido. El nectario anular parece ser un carácter compartido por toda la subtribu Psittacanthinae, que lo diferencia de la subtribu Ligarinae con nectario estilar. El gineceo unilocular formado por una estructura central única es un carácter compartido con otros géneros de la subtribu Psittacanthinae, con la excepción de Tripodanthus. El androceo está formado por estambres bitecados, tetrasporangiados con lóculos septados que aquí se consideran una adaptación para liberar polen durante un período prolongado de tiempo, en lugar de sugerencias previas que lo explican como resultado de la presión evolutiva para reducir el tamaño de la antera o para facilitar la nutrición de microesporas en anteras grandes.

Tracking Host Trees: The Phylogeography of Endemic Psittacanthus sonorae (Loranthaceae) Mistletoe in the Sonoran Desert.

The host dependence of mistletoes suggests that they track the distributions of their hosts. However, the factors that determine the geographic distribution of mistletoes are not well understood. In this study, the phylogeography of Psittacanthus sonorae was reconstructed by sequencing one nuclear (ITS) and two plastid (trnL-F and atpB-rbcL) regions of 148 plants from populations separated by the Sea of Cortez. Divergence time and gene flow were estimated to gain insight into the historical demography and geographic structuring of genetic variation. We also described and mapped the spatial distribution of suitable habitat occupied by P. sonorae and its most common host Bursera microphylla in the Sonoran Desert, along with their responses to Quaternary climate fluctuations using environmental data and ecological niche modeling (ENM). We detected environmental and genetic differentiation between the peninsular and continental P. sonorae populations. Population divergence occurred during the Pleistocene, around the time of the Last Glacial Maximum. No signals of population growth were detected, with net gene flow moving from the continent to the peninsula. ENM models indicate decoupled responses by the mistletoe and its main host to past climate changes. For the Last Interglacial to the present, most models produce only partial areas of overlap on both the peninsula and the continent. Our results support a scenario of Late-Pleistocene isolation and divergence with asymmetrical gene flow between peninsular and continental P. sonorae populations. Continental populations migrated to the peninsula and the spatial isolation probably produced genetic differentiation under different environmental conditions.

Historical biogeography of Loranthaceae (Santalales): Diversification agrees with emergence of tropical forests and radiation of songbirds.

Coadaptation between mistletoes and birds captured the attention of Charles Darwin over 150 years ago, stimulating considerable scientific research. Here we used Loranthaceae, a speciose and ecologically important mistletoe family, to obtain new insights into the interrelationships among its hosts and dispersers. Phylogenetic analyses of Loranthaceae were based on a dataset of nuclear and chloroplast DNA sequences. Divergence time estimation, ancestral area reconstruction, and diversification rate analyses were employed to examine historical biogeography. The crown group of Loranthaceae was estimated to originate in Australasian Gondwana during the Paleocene to early Eocene (59 Ma, 95% HPD: 53-66 Ma), and rapidly diversified, converting from root parasitic to aerial parasitic trophic mode ca. 50 Ma during the Eocene climatic optimum. Subsequently, Loranthaceae were inferred to be widespread in Australasia and South America but absent in Africa. The African and European members were derived from Asiatic lineages. The burst of diversification of Loranthaceae occurred during a climatic optimum period that coincides with the dominance of tropical forests in the world. This also corresponds to the trophic mode conversion of Loranthaceae and rapid radiation of many bird families - important agents for long-distance dispersal in the Cenozoic.

Comparative inflorescence development in selected Andean Santalales.

PREMISE OF THE STUDY: Loranthaceae, Santalaceae, and Viscaceae are the most diversified hemiparasitic families of Santalales in the Andes. Their partial inflorescences (PIs) vary from solitary flowers, or dichasia in most Santalales, to congested floral groups along articles in most Viscaceae. The atypical articled inflorescences in Phoradendreae (Viscaceae), a phylogenetic novelty restricted to this tribe, have been variously described as racemes, spikes, fascicles, or as intercalary inflorescences, but no developmental studies have been performed to compare them with the construction of PIs across Santalales. METHODS: We used standard light microscopy and scanning electron microscopy to record the inflorescence development in members of Phoradendreae (Viscaceae) in comparison to those in species of Aetanthus, Gaiadendron, Oryctanthus, Passovia, and Peristethium (Loranthaceae) and Antidaphne (Santalaceae). KEY RESULTS: Morphological and developmental comparisons as well as optimization onto a phylogenetic framework indicate that individual inflorescences in Santalales are indeterminate and are formed by axillary cymose PIs. The latter correspond to dichasia, either simple, compound, or variously reduced by abortion of lateral flowers, abortion of the terminal flower, or loss of bracteoles. CONCLUSIONS: Dichasia are plesiomorphic in Santalales. These results favor the interpretation that inflorescences in Phoradendreae are formed by the fusion of serial dichasia (=floral rows) with the main inflorescence axis via syndesmy. We compared this interpretation with the competing one based on the co-occurrence of collateral and serial floral buds.

A mistletoe tale: postglacial invasion of Psittacanthus schiedeanus (Loranthaceae) to Mesoamerican cloud forests revealed by molecular data and species distribution modeling.

BACKGROUND: Ecological adaptation to host taxa is thought to result in mistletoe speciation via race formation. However, historical and ecological factors could also contribute to explain genetic structuring particularly when mistletoe host races are distributed allopatrically. Using sequence data from nuclear (ITS) and chloroplast (trnL-F) DNA, we investigate the genetic differentiation of 31 Psittacanthus schiedeanus (Loranthaceae) populations across the Mesoamerican species range. We conducted phylogenetic, population and spatial genetic analyses on 274 individuals of P. schiedeanus to gain insight of the evolutionary history of these populations. Species distribution modeling, isolation with migration and Bayesian inference methods were used to infer the evolutionary transition of mistletoe invasion, in which evolutionary scenarios were compared through posterior probabilities. RESULTS: Our analyses revealed shallow levels of population structure with three genetic groups present across the sample area. Nine haplotypes were identified after sequencing the trnL-F intergenic spacer. These haplotypes showed phylogeographic structure, with three groups with restricted gene flow corresponding to the distribution of individuals/populations separated by habitat (cloud forest localities from San Luis Potosí to northwestern Oaxaca and Chiapas, localities with xeric vegetation in central Oaxaca, and localities with tropical deciduous forests in Chiapas), with post-glacial population expansions and potentially corresponding to post-glacial invasion types. Similarly, 44 ITS ribotypes suggest phylogeographic structure, despite the fact that most frequent ribotypes are widespread indicating effective nuclear gene flow via pollen. Gene flow estimates, a significant genetic signal of demographic expansion, and range shifts under past climatic conditions predicted by species distribution modeling suggest post-glacial invasion of P. schiedeanus mistletoes to cloud forests. However, Approximate Bayesian Computation (ABC) analyses strongly supported a scenario of simultaneous divergence among the three groups isolated recently. CONCLUSIONS: Our results provide support for the predominant role of isolation and environmental factors in driving genetic differentiation of Mesoamerican parrot-flower mistletoes. The ABC results are consistent with a scenario of post-glacial mistletoe invasion, independent of host identity, and that habitat types recently isolated P. schiedeanus populations, accumulating slight phenotypic differences among genetic groups due to recent migration across habitats. Under this scenario, climatic fluctuations throughout the Pleistocene would have altered the distribution of suitable habitat for mistletoes throughout Mesoamerica leading to variation in population continuity and isolation. Our findings add to an understanding of the role of recent isolation and colonization in shaping cloud forest communities in the region.

Anatomía Floral de Peristethium leptostachyum (Loranthaceae) / Floral anatomy of Peristethium leptostachyum (Loranthaceae)

ResumenPeristethium leptostachyum es una especie hemiparásita de la familia Loranthaceae, distribuida en Colombia, Costa Rica, Ecuador, Perú, Venezuela y Panamá. Previamente tratada como Struthanthus leptostachyus, la especie fuerecientemente fue reubicada en Peristethium junto con otras que previamente estaban en los géneros Cladocolea y Struthanthus. La decisión de reconocer a Peristethium como género es controversial y fue tomada con base en caracteres de la inflorescencia y de la flor; en tanto que la monofilia de los tres géneros nombrados es incierta. En esta investigación se estudió la morfoanatomía de flores e inflorescencias de Peristethium leptostachyum, detallando la estructura del androceo y gineceo, así como los procesos de microgametogénesis y megagametogénesis; adicionalmente se realizaron comparaciones con especies afines y precisiones en relación con las diagnosis previas. Se recolectaron flores en diversas fases de desarrollo en Santa María (Boyacá-Colombia), se prepararon y analizaron bajo microscopio secciones histológicas teñidas con astrabluefucsina, además de disecciones bajo estereomicroscopio. Los resultados mostraron que P. leptostachyum comparte caracteres inflorenciales con Cladocolea (inflorescencia determinada, flor terminal ebracteada), pero también con Struthanthus (pares de tríadas a lo largo del eje, brácteas caducas y flores actinomorfas). Las flores de P. Leptostachyum de Santa María son claramente hermafroditas, con androceos y gineceos totalmente desarrollados; lo cual contradice la descripción hecha por Kuijt que reporta una condición dioica para esta especie. El androceo resultó afín al de Struthanthus vulgaris, con tapetum glandular y microsporogénesis simultánea; en contraste, Cladocolea loniceroides presenta tapetum periplasmodial y microesporogénesis sucesiva. El gineceo de P. leptostachyum, al igual que en Cladocolea, Struthanthus y Phthirusa, es unilocular con mamelón y tejido arquesporial orientado hacia el estilo, el cual es sólido y con tejido amilífero. P. leptostachyum es afín a Cladocolea loniceroides y difiere de Struthanthus vulgaris por presentar varios sacos embrionarios y pelvis (hipostasa) no lignificada. La presencia de un canal estilar sólido se propone como sinapomorfía de la tribu Psittacanthinae. Dado que P. Leptostachyum comparte caracteres anatómicos florales tanto con Cladocolea como con Strutanthus, la relación entre estos tres géneros no queda resuelta, se requieren estudios filogenéticos para establecer esta relación y poner a prueba las hipótesis de monofilia de cada uno de ellos.

AbstractPeristethium leptostachyum is a hemiparasite species of the family Loranthaceae, distributed in Colombia, Costa Rica, Ecuador, Peru, Venezuela and Panama. Previously treated as Struthanthus leptostachyus, the species was recently transferred to Peristethium together with other species of Cladocolea and Struthanthus. The present research describes the inflorescence and floral morphoanatomy of Peristethium leptostachyum, detailing the structure of the androecium and gynoecium and the processes of microgametogenesis and megagametogenesis, thus allowing comparison with Struthanthus and Cladocolea. Flowering material was collected in February and August 2012, in Santa María, Boyacá, Colombia. Histological sections were prepared and stained with astrablue-fuchsin and floral dissections were performed under a stereomicroscope. Peristethium leptostachyum shares inflorescence characters with Cladocolea (determinate inflorescence, ebracteate terminal flower), but also with Struthanthus (pairs of triads along the axis, deciduous bracts and actinomorphic flowers). The flowers of P. leptostachyum from Santa María are clearly hermaphrodites with androecium and gynoecium fully developed. This observation contradicts the description by Kuijt who reported this species to be dioecious. The androecium was observed to be similar to that of Struthanthus vulgaris, with a glandular tapetum and simultaneous microsporogenesis; in contrast, Cladocolea loniceroides has a periplasmodial tapetum and successive microsporogenesis. The gynoecium of P. leptostachyum, like that of Cladocolea, Struthanthus and Phthirusa, has a unilocular ovary with a mamelon and arquesporial tissue isoriented towards the style, which in turn is solid and amyliferous. Peristethium leptostachyum is similar to Cladocolea loniceroides and differs from Strutanthus vulgaris in presenting multiple embryo sacs and an unlignified pelvis (hipostase). The presence of a solid stylar canal is proposed as a synapomorphy of the tribe Psittacanthinae. Given that P. leptostachyum shares characters with both Cladocolea and Struthanthus generic placement cannot be clearly determined on the basis of anatomical evidence. Phylogenetic studies that include representative species of all three genera are desirable to test hypotheses of monophyly. The sexual system observed here in P. leptostachyum is different from that reported by Kuijt and more studies are needed to identify the factors (geographic, ecological, etc.) that influence this variation. Rev. Biol. Trop. 64 (1): 341-352. Epub 2016 March 01.

[Floral anatomy of Peristethium leptostachyum (Loranthaceae)]. / Anatomía Floral de Peristethium leptostachyum (Loranthaceae).

Peristethium leptostachyum is a hemiparasite species of the family Loranthaceae, distributed in Colombia, Costa Rica, Ecuador, Peru, Venezuela and Panama. Previously treated as Struthanthus leptostachyus, the species was recently transferred to Peristethium together with other species of Cladocolea and Struthanthus. The present research describes the inflorescence and floral morphoanatomy of Peristethium leptostachyum, detailing the structure of the androecium and gynoecium and the processes of microgametogenesis and megagametogenesis, thus allowing comparison with Struthanthus and Cladocolea. Flowering material was collected in February and August 2012, in Santa María, Boyacá, Colombia. Histological sections were prepared and stained with astrablue-fuchsin and floral dissections were performed under a stereomicroscope. Peristethium leptostachyum shares inflorescence characters with Cladocolea (determinate inflorescence, ebracteate terminal flower), but also with Struthanthus (pairs of triads along the axis, deciduous bracts and actinomorphic flowers). The flowers of P. leptostachyum from Santa María are clearly hermaphrodites with androecium and gynoecium fully developed. This observation contradicts the description by Kuijt who reported this species to be dioecious. The androecium was observed to be similar to that of Struthanthus vulgaris, with a glandular tapetum and simultaneous microsporogenesis; in contrast, Cladocolea loniceroides has a periplasmodial tapetum and successive microsporogenesis. The gynoecium of P. leptostachyum, like that of Cladocolea, Struthanthus and Phthirusa, has a unilocular ovary with a mamelon and arquesporial tissue isoriented towards the style, which in turn is solid and amyliferous. Peristethium leptostachyum is similar to Cladocolea loniceroides and differs from Strutanthus vulgaris in presenting multiple embryo sacs and an unlignified pelvis (hipostase). The presence of a solid stylar canal is proposed as a synapomorphy of the tribe Psittacanthinae. Given that P. leptostachyum shares characters with both Cladocolea and Struthanthus generic placement cannot be clearly determined on the basis of anatomical evidence. Phylogenetic studies that include representative species of all three genera are desirable to test hypotheses of monophyly. The sexual system observed here in P. leptostachyum is different from that reported by Kuijt and more studies are needed to identify the factors (geographic, ecological, etc.) that influence this variation.

Flora of the State of Paraíba, Brazil: Loranthaceae Juss / Flora do Estado da Paraíba, Brasil: Loranthaceae Juss

The family Loranthaceae Juss., characterized by its parasitic habit, is rarely studied in Brazil. Current research provides a taxonomic survey of Loranthaceae in the State of Paraiba, northeastern Brazil, in which ten species belonging to four genera have been recorded: Passovia (one species), Psittacanthus (two species), Pusillanthus (one species) and Struthanthus (six species). Struthanthus concinnus Mart. was found for the first time in the state of Paraíba. A key to the species identification and taxonomic descriptions is presented, with images, geographic distribution and host data.

A família Loranthaceae Juss. é caracterizada pelo hábito parasítico e é ainda escassamente estudada no Brasil. Este trabalho realizou o levantamento taxonômico de Loranthaceae no Estado da Paraíba, nordeste do Brasil, onde dez espécies pertencentes a quatro gêneros foram registradas: Passovia (1 espécie), Psittacanthus (2 espécies), Pusillanthus (1 espécie) e Struthanthus (6 espécies). Struthanthus concinnus Mart. foi registrada pela primeira vez para a flora do Estado da Paraíba. Uma chave para a identificação das espécies e descrições taxonômicas é apresentada, com imagens, dados de distribuição geográfica e de hospedeiros também fornecidos.

Medicinal parasitic plants on diverse hosts with their usages and barcodes.

Medicinal properties of parasitic plants were investigated by means of ethnobotanical study in some areas of northeastern Thailand. Important traditional usages are: Scurrula atropurpurea nourishes blood, Dendrophthoe pentandra decreases high blood pressure, and Helixanthera parasitica treats liver disease. Their systematics were also determined. The research is based on findings obtained from 100 parasite-host pairs. Of these, eight parasitic species were recorded; they are members of two families, viz. family Loranthaceae, namely D. lanosa, D. pentandra, H. parasitica, Macrosolen brandisianus, M. cochinchinensis and S. atropurpurea, and family Viscaceae, namely Viscum articulatum and V. ovalifolium. In addition, each parasitic species is found on diverse hosts, indicating non-host-parasitic specificity. Species-specific tagging of all species studied was carried out using the rbcL and psbA-trnH chloroplast regions. These tag sequences are submitted to GenBank databases under accession numbers JN687563-JN687578. Genetic distances calculated from nucleotide variations in a couple of species of each genus, Dendrophthoe, Macrosolen, and Viscum, were 0.032, 0.067 and 0.036 in the rbcL region, and 0.269, 0.073 and 0.264 in the psbA-trnH spacer region, respectively. These variations will be used for further identification of incomplete plant parts or other forms such as capsule, powder, dried or chopped pieces.

Occurrence of the lutein-epoxide cycle in mistletoes of the Loranthaceae and Viscaceae.

The lutein-epoxide cycle (Lx cycle) is an auxiliary xanthophyll cycle known to operate only in some higher-plant species. It occurs in parallel with the common violaxanthin cycle (V cycle) and involves the same epoxidation and de-epoxidation reactions as in the V cycle. In this study, the occurrence of the Lx cycle was investigated in the two major families of mistletoe, the Loranthaceae and the Viscaceae. In an attempt to find the limiting factor(s) for the occurrence of the Lx cycle, pigment profiles of mistletoes with and without the Lx cycle were compared. The availability of lutein as a substrate for the zeaxanthin epoxidase appeared not to be critical. This was supported by the absence of the Lx cycle in the transgenic Arabidopsis plant lutOE, in which synthesis of lutein was increased at the expense of V by overexpression of epsilon-cyclase, a key enzyme for lutein synthesis. Furthermore, analysis of pigment distribution within the mistletoe thylakoids excluded the possibility of different localizations for the Lx- and V-cycle pigments. From these findings, together with previous reports on the substrate specificity of the two enzymes in the V cycle, we propose that mutation to zeaxanthin epoxidase could have resulted in altered regulation and/or substrate specificity of the enzyme that gave rise to the parallel operation of two xanthophyll cycles in some plants. The distribution pattern of Lx in the mistletoe phylogeny inferred from 18S rRNA gene sequences also suggested that the occurrence of the Lx cycle is determined genetically. Possible molecular evolutionary processes that may have led to the operation of the Lx cycle in some mistletoes are discussed.