Article
To read the full-text of this research, you can request a copy directly from the authors.

Abstract

The Boraginales are now universally accepted as monophyletic and firmly placed in Lamiidae. However, a consensus about familial classification has remained elusive, with some advocating recognition of a single, widely variable family, and others proposing recognition of several distinct families. A consensus classification is proposed here, based on recent molecular phylogenetic studies, morphological characters, and taking nomenclatural stability into consideration. We suggest the recognition of eleven, morphologically well-defined and clearly monophyletic families, namely the Boraginaceae s.str., Codonaceae, Coldeniaceae fam. nov., Cordiaceae, Ehretiaceae, Heliotropiaceae, Hoplestigmataceae, Hydrophyllaceae, Lennoaceae, Namaceae, and Wellstediaceae. Descriptions, synonomy, a taxonomic key, and a list of genera for these eleven families are provided, including the new family Coldeniaceae (monogeneric) and Namaceae (segregated from Hydrophyllaceae and comprising Nama, Eriodictyon, Turricula, and Wigandia), the latter necessitating a revised circumscription of a more morphologically coherent Hydrophyllaceae.

No full-text available

Request Full-text Paper PDF

To read the full-text of this research,
you can request a copy directly from the authors.

... Euploca Nutt. (=Heliotropium L. section Orthostachys R.Br.) (Boraginaceae sensu APG IV or Heliotropiaceae) is a widespread genus with c. 100-120 species (Frohlich, 1978;Birecka, Frohlich & Glickman, 1983;Luebert et al., 2016). It exhibits remarkable variation in physiological, morphological, chemical, reproductive and life-history features, including various intermediate states, making it a potentially valuable system for evolutionary studies. ...
... Molecular studies have demonstrated that Hydrophyllaceae and Lennoaceae fall within the broader concepts of Boraginaceae, resulting in the expanded Boraginaceae of APG III and APG IV (APG III, 2009;Reveal & Chase, 2011;APG IV, 2016). To render families more homogeneous, Luebert et al. (2016) divided Boraginaceae sensu APG into 11 narrower families, and placed in Heliotropiaceae all the species of Heliotropium and Tournefortia, as traditionally understood, essentially from the time of Linnaeus (1753: 130, 140) through to the end of the 20 th century: traditionally, Heliotropium and Tournefortia were distinguished by the key character, respectively, of dry versus moist fruit. Here we refer to those traditional circumscriptions as Heliotropium s.l. and Tournefortia s.l., although the designation 's.l.' for the former may be problematic, as noted below. ...
... For example, the inflorescences of E. tenella (clade 4) have a gross aspect so similar to the vegetative shoots that one may easily overlook some inflorescences when observing the whole plant (Fig. 14A). Such inflorescences are termed anthoclades (sensu Luebert et al., 2016). By contrast, in the bractless species, inflorescence axes are typically significantly thinner than vegetative shoots, with flowers much more closely spaced than are the leaves of vegetative shoots (Fig. 14B). ...
Article
Full-text available
We present a phylogenetic analysis using plastid (matK, rbcL) and nuclear (nrITS) DNA for diverse Euploca spp. (formerly Heliotropium section Orthostachys) from the worldwide distribution of a genus and including species encompassing the wide physiological and morphological diversity of the genus. Our results indicate that some remarkably complex features arose multiple times in parallel in Euploca, including attributes of its subsections under section Orthostachys, notably plants that, above ground, consist almost entirely of inflorescences. To elucidate in greater detail the distribution of C4 species in Euploca and Heliotropium s.s., we made > 800 δ 13C determinations, including some from the traditional genus Tournefortia. We greatly increase the number of proven C4 species in Euploca, but found none outside Euploca. Of the tested Euploca spp., c. 28% are C3 or intermediate in carbon fixation pathway. Our phylogenetic results indicate four parallel/convergent acquisitions of C4 photosynthesis or fewer origins with subsequent loss in some species.
... Hydrophyllaceae and Namaceae are successively sister families to the remainder of the Boraginales II clade (including also Coldeniaceae, Cordiaceae, Ehretiaceae, Heliotropiaceae, Hoplestigmataceae, and Lennoaceae) based on plastid data (Luebert et al., 2016;Stull et al., 2015;Weigend et al., 2014). Phylogenetic studies over the last two decades have resolved the relationships in these two to families: Hydrophyllaceae falls into three major clades (Ferguson, 1998;Walden, 2010). ...
... ex S. Watson, Howellanthus (Constance) Walden & R. Patt. and Hesperochiron S. Watson (unnamed;Luebert et al., 2016;Walden et al., 2014). Namaceae falls into two clades: a Nama clade (about 50 spp.; ...
... Hofmann et al., 2016), and a clade comprising woody Wigandia Kunth (6 spp.; Hofmann et al., 2016), Eriodictyon Benth., and Turricula J.F. Macbr. plus two species of Nama (rendering Nama polyphyletic Ferguson, 1998;Luebert et al., 2016;Taylor, 2012). ...
Article
This study aimed to examine the systematic position of South American species of Phacelia (Hydrophyllaceae) and Wigandia (Namaceae) and the historical biogeography of Hydrophyllaceae and Namaceae using molecular dating and ancestral area reconstruction. To this end, we constructed two datasets, one with a plastid (ndhF) and one with a nuclear marker (ITS), using previously published and newly generated sequences. We inferred the phylogeny of Hydrophyllaceae and Namaceae implementing both likelihood and Bayesian methods. We also estimated divergence times and ancestral areas for all major clades using a relaxed Bayesian uncorrelated molecular clock and the dispersal-extinction-cladogenesis (DEC) approach, respectively. The South American representatives of Phacelia are placed in three different clades of the genus and two colonizations of South America by North American species took place in the Miocene and at least one in the late Pliocene. Wigandia forms a well-supported monophylum with interspecific relationships partly unresolved. Within Namaceae a colonization of South America by North American species occurred during the Oligocene-Miocene transition. The MRCA of Wigandia was distributed in North and Central America in late Oligocene. Long-distance dispersal may have been necessary for the colonization of South America by Namaceae during the Oligocene-Miocene transition, when North and South America were not connected.
... Heliotropiaceae was segregated from Boraginaceae s.l. (BWG 2016) and comprises approximately 450 species belonging to four genera, Euploca Nuttall, Heliotropium L., Myriopus Small, and the monotypic Ixorhea Fenzl, endemic from Argentina (BWG 2016). This family is widely distributed, although it is mainly concentrated in tropical and subtropical regions (Diane et al. 2016). ...
... Heliotropiaceae was segregated from Boraginaceae s.l. (BWG 2016) and comprises approximately 450 species belonging to four genera, Euploca Nuttall, Heliotropium L., Myriopus Small, and the monotypic Ixorhea Fenzl, endemic from Argentina (BWG 2016). This family is widely distributed, although it is mainly concentrated in tropical and subtropical regions (Diane et al. 2016). ...
... Representatives of Heliotropiaceae are herbs to trees with simple and alternate leaves; pentamerous flowers, bisexual, solitary or arranged in terminal or axillary inflorescences of the thyrsus or scorpioids type; gynoecium frequently with nectariferous disk at the base of the ovary; fleshy or dry fruit, usually with 4 (rare 1-2) seeds. The family synapomorphies encompass the presence of a terminal style with a conical stigmatic head and the ring-shaped basal stigma (BWG 2016). ...
Article
Full-text available
The taxonomic floristic survey of the Heliotropiaceae species in Ceará state is part of the “Flora of Ceará: knowing to conserve” project. The study was based on specimens deposited in national and international herbaria, relevant bibliography, images of type specimens, and field trips. Fourteen species belonging to three genera were recorded: Euploca (six spp.) was the most representative in number of species, followed by Heliotropium and Myriopus (four spp. each). Euploca paradoxa is endemic to Brazil, being a new occurrence for Ceará. Most species occur in Conservation Units, except for Euploca humilis, E. paradoxa, Heliotropium angiospermum, and H. funkiae. The species occur predominantly in dry habitats, such as Stepic Savanna (Caatinga), however some species have been recorded in more humid areas, such as Ombrophilous Forest and coastal vegetation.
... Pantropical Cordiaceae are an integral element of Boraginales and comprise some 400 species of trees and shrubs Luebert et al. 2016). They are widespread in the Americas, Africa and Asia, with the greatest diversity in tropical America, and are mostly found in seasonally dry areas and permanently humid habitats (Gürke 1893;Miller 1985). ...
... Varronia differs from Cordia in its general habit (multi-stemmed shrubs and lianas versus mostly trees), leaf morphology (serrate versus entire margin), inflorescence architecture (syndesmia, with the paracladia entirely incorporated in the inflorescence axis, Uhlarz and Weberling 1977, versus acropetal, corymbose thyrses) and pollen morphology (porate versus colporate apertures; Miller and Gottschling 2007;Gottschling et al. 2016). Molecular phylogenetics (Gottschling et al. 2005;Weigend et al. 2014) further show the close relationship between Cordiaceae, monotypic Coldeniaceae (Coldenia procumbens L.) Table 1 Comparison between the two species under investigation and to other Boraginales (more traits are tabulated in, e.g., Khaleel 1985 andLuebert et al. 2016 and Hoplestigmataceae (Hoplestigma Pierre, comprising two species). However, the identified relationships still await corroboration from morphological data of these poorly investigated taxa. ...
... Cordiaceae belong to a plant group that has been denoted as Primary Woody Boraginales (PWB, i.e., Cordiaceae plus Coldeniaceae and Hoplestigmataceae, Ehretiaceae plus Lennoaceae, Heliotropiaceae;Gottschling 2003;Luebert et al. 2016), in generative aspects being characterised by the presence of a lignified and multi-layered endocarp (Table 1). Many PWB have drupaceous fruits (or derivatives thereof), and the endocarp protects the embryo during the passage of the digestive tract of animals or during seasonally unfavourable ecological conditions. ...
Article
Despite their ecological importance and wide distribution, Cordiaceae have not been subject to detailed anatomical study yet. We examined flower and fruit anatomies of Cordia nodosa and Varronia bonplandii (using paraffin sectioning and light microscopy) in comparison with other woody members of Boraginales. The internal architecture of the superior bicarpellate ovary resulted from the development of secondary septa including apical, basal and false septa, as reported also from other Boraginales. Novel characters of Cordiaceae were identified such as an extensive tissue surrounding the gynoecium of C. nodosa and a lignified cap on top of the endocarp in V. bonplandii. Fruits containing a single seed may have originated several times independently in Cordiaceae deriving from the four-seeded condition present today in most other non-capsular Boraginales. Distyly of V. bonplandii does not appear genetically fixed but may result from different ecological conditions. Hemianatropous ovules are now considered the derived character state in Cordiaceae, originating from anatropous ovules present in other Boraginales. Studies like this improve the knowledge about generative organs in general and identify phylogenetically informative characters. However, better knowledge of functional morphology and ecological importance regarding many traits identified require further investigations on the diversity of Cordiaceae.
... The sampling here includes four additional families, Codonaceae, Namaceae, Cordiaceae, and Coldeniaceae, compared with the 1KP study. Our analyses indicate that Boraginales contain two suprafamilial clades: clade I consisting of Codonaceae þ Boraginaceae with 100% BS in all six trees (Boraginales I, Weigend et al. 2014), and clade II, including the remaining seven Boraginales families with maximal BS support ( fig. 4) Stull et al. (2015), and Luebert et al. (2016), which supported Hydrophyllaceae and Namaceae as successive sisters to the remaining families. Conversely, recently published plastid phylogenetics and morphological character analyses support the hypothesis that Heliotropiaceae are sister to a well-supported clade consisting of two clades, one including Cordiaceae, Hoplestigmataceae, and Coldeniaceae and the other including Ehretiaceae and Lennoaceae, collectively form a super clade with species bearing multilayered endocarp and four or fewer ovules (Refulio-Rodriguez and Olmstead 2014; Weigend et al. 2014;Luebert et al. 2016). ...
... Our analyses indicate that Boraginales contain two suprafamilial clades: clade I consisting of Codonaceae þ Boraginaceae with 100% BS in all six trees (Boraginales I, Weigend et al. 2014), and clade II, including the remaining seven Boraginales families with maximal BS support ( fig. 4) Stull et al. (2015), and Luebert et al. (2016), which supported Hydrophyllaceae and Namaceae as successive sisters to the remaining families. Conversely, recently published plastid phylogenetics and morphological character analyses support the hypothesis that Heliotropiaceae are sister to a well-supported clade consisting of two clades, one including Cordiaceae, Hoplestigmataceae, and Coldeniaceae and the other including Ehretiaceae and Lennoaceae, collectively form a super clade with species bearing multilayered endocarp and four or fewer ovules (Refulio-Rodriguez and Olmstead 2014; Weigend et al. 2014;Luebert et al. 2016). The relationships among Boraginales families are also supported by previous fruit morphological and phylogenetic analyses (Weigend et al. 2014;Luebert et al. 2016). ...
... Conversely, recently published plastid phylogenetics and morphological character analyses support the hypothesis that Heliotropiaceae are sister to a well-supported clade consisting of two clades, one including Cordiaceae, Hoplestigmataceae, and Coldeniaceae and the other including Ehretiaceae and Lennoaceae, collectively form a super clade with species bearing multilayered endocarp and four or fewer ovules (Refulio-Rodriguez and Olmstead 2014; Weigend et al. 2014;Luebert et al. 2016). The relationships among Boraginales families are also supported by previous fruit morphological and phylogenetic analyses (Weigend et al. 2014;Luebert et al. 2016). ...
Article
Full-text available
Asterids are one of the most successful angiosperm lineages, exhibiting extensive morphological diversity and including a number of important crops. Despite their biological prominence and value to humans, the deep asterid phylogeny has not been fully resolved, and the evolutionary landscape underlying their radiation remains unknown. To resolve the asterid phylogeny, we sequenced 213 transcriptomes/genomes and combined them with other datasets, representing all accepted orders and nearly all families of asterids. We show fully supported monophyly of asterids, Berberidopsidales as sister to asterids, monophyly of all orders except Icacinales, Aquifoliales and Bruniales, and monophyly of all families except Icacinaceae and Ehretiaceae. Novel taxon placements benefited from the expanded sampling with living collections from botanical gardens, resolving hitherto uncertain relationships. The remaining ambiguous placements here are likely due to limited sampling and could be addressed in the future with relevant additional taxa. Using our well-resolved phylogeny as reference, divergence time estimates support an Aptian (Early Cretaceous) origin of asterids and the origin of all orders before the K-Pg boundary. Ancestral state reconstruction at the family level suggests that the asterid ancestor was a woody terrestrial plant with simple leaves, bisexual and actinomorphic flowers with free petals and free anthers, a superior ovary with a style, and drupaceous fruits. WGD analyses provide strong evidence for 33 WGDs in asterids and one in Berberidopsidales, including four supra-familial and seven familial/sub-familial WGDs. Our results advance the understanding of asterid phylogeny and provide numerous novel evolutionary insights into their diversification and morphological evolution.
... Many old sources and descriptions labeled "Boraginaceae" deal with a broader sense of the family concept. First described by Jussieu (1789), the borage family has been variously circumscribed, ranging from an inclusive taxon encompassing the whole order Boraginales to a family defined in its most strict sense by the Boraginales Working Group (Luebert et al., 2016), with around 90 genera (see Appendix 2 in Luebert et al. [2016]) and some 1600 to 1700 species. The recent classification of Boraginales as outlined by Luebert et al. (2016) reflects current knowledge about phylogenetic relationships in the clade and provides a familial classification for the order. ...
... Many old sources and descriptions labeled "Boraginaceae" deal with a broader sense of the family concept. First described by Jussieu (1789), the borage family has been variously circumscribed, ranging from an inclusive taxon encompassing the whole order Boraginales to a family defined in its most strict sense by the Boraginales Working Group (Luebert et al., 2016), with around 90 genera (see Appendix 2 in Luebert et al. [2016]) and some 1600 to 1700 species. The recent classification of Boraginales as outlined by Luebert et al. (2016) reflects current knowledge about phylogenetic relationships in the clade and provides a familial classification for the order. ...
... First described by Jussieu (1789), the borage family has been variously circumscribed, ranging from an inclusive taxon encompassing the whole order Boraginales to a family defined in its most strict sense by the Boraginales Working Group (Luebert et al., 2016), with around 90 genera (see Appendix 2 in Luebert et al. [2016]) and some 1600 to 1700 species. The recent classification of Boraginales as outlined by Luebert et al. (2016) reflects current knowledge about phylogenetic relationships in the clade and provides a familial classification for the order. In accordance with this scheme, Boraginaceae s. str. is a widely distributed group in tropical, subtropical, and temperate regions. ...
Article
During the preparation of the treatment of the family Boraginaceae s. str. for Argentina, dozens of names were identified as needing typifications to stabilize their nomenclature. As a result, lectotypes are designated for 11 names (Allocarya alternifolia Brand, Amsinckia angustifolia Lehm. var. microcarpa Speg., Amsinckia tessellata A. Gray, Echinospermum patagonicum Speg., Eritrichium albiflorum Griseb., Eritrichium humile (Ruiz & Pav.) DC. var. capillatum Clos, Eritrichium humile (Ruiz & Pav.) DC. var. congestum Wedd., Gruvelia pusilla A. DC., Lithospermum chilense Colla, Myosotis corymbosa Ruiz & Pav., and Pectocarya chilensis DC.). Additionally, second-step lectotypes are designated for nine names in Eritrichium Schrad. ex Gaudin (E. calandrinioides Phil., E. cinereum Phil., E. flavicans Phil., E. germainii Phil., E. graminifolium Phil., E. pedicellare Phil., E. pratense Phil., E. pugae Phil., and E. pulchellum Phil.).
... We first calculated net diversification rates at crown nodes of major clades of Lithospermeae using the R-package GEIGER 2.0.3 (Harmon et al., 2008). Species richness of each clade was sourced from Luebert et al. (2016) and Weigend et al. (2016). In order to evaluate whether diversification rates are significantly high across the Lithospermeae clades, as compared to its background diversification rate, we built 95% confidence intervals (CI) for crown-and stem node-diversification rates. ...
... This model is appropriate in the case of Lithospermeae, because several clades in the group appear to have originated after a long-distance dispersal event (Chacón et al., 2017), similarly to other Boraginaceae clades such as Cynoglossoideae and Omphalodeae (Otero et al., 2019a, b). Seven areas were defined, representing the entire distribution range of Lithospermeae, using occurrence data from specimens sequenced in this study, as well as floras and taxonomic revisions by Brand (1930), Johnston (1952Johnston ( , 1953, Ball (1972), Al-Shehbaz (1991), Böhle et al. (1996), Hilger and Böhle (2000), Boyd (2002Boyd ( , 2004, Cecchi and Selvi (2009), Buys (2011), Cecchi et al. (2011, Coppi et al. (2015), Weigend et al. (2009Weigend et al. ( , 2016, Luebert et al. (2016), and Cohen (2018). Areas were coded as A: South America, where five species of Lithospermum occur. ...
Article
Studies about the drivers of angiosperm clade diversifications have revealed how the environment continuously alters the species chances to adapt or to go extinct. This process depends on complex interactions between abiotic and biotic factors, conditioned to the geological and tectonic settings, the genetic variability of species and the rate at which speciation occurs. In this study, we aim to elucidate the timing of diversification of the Lithospermeae, the second largest tribe within Boraginaceae, and to identify the possible morphological and ecological characters associated with shifts in diversification rates of the most species-rich clades. Lithospermeae includes ca. 470 species and 26 genera, among which are some of the largest genera of the family such as Onosma (150 spp.), Echium (60 spp.), and Lithospermum (80 spp.). An exhaustive study of the whole clade is not available to date and its evolutionary history and diversification rates are incompletely known. In the present study, we provide the most comprehensive phylogeny of the group so far, sampling 242 species and all 26 genera. We found that crown-groups and diversification rates of Lithospermeae largely date back to the Mid-Miocene, with high diversification rates in the largest genera, though only significantly high in Onosma. Our analysis fails to associate any of the functional or morphological traits considered with significant shifts in diversification rates. The timing of the diversification of the species-rich clades corresponds with Miocene tectonic events and global climate changes increasing aridity across Eurasia and western North America. These results suggest a causal link between known ecological features of Lithospermeae (i.e., pre-adaptation to arid, open habitats, and mineral soils) and their diversification. Future studies should expand the sampling of individual subclades and detailed functional analyses to identify the contribution of adaptations to arid conditions and pollinator shifts.
... Currently, these subfamilies are largely recognized at the family level. Recently, Luebert et al. (2016) suggested the recognition of 11 monophyletic families: Boraginaceae s.s., Codonaceae, Coldeniaceae, Cordiaceae, Ehretiaceae, Heliotropiaceae, Hoplestigmataceae, Hydrophyllaceae, Lennoaceae, Namaceae and Wellstediaceae. ...
... In the present study, Boraginaceae are used in the strict sense (Boraginaceae s.s.; see Luebert et al., 2016), and the family currently includes c. 90 genera and c. 1600 species distributed across three subfamilies: Echiochiloideae, Boraginoideae and Cynoglossoideae (Chac on et al., 2016). Boraginoideae and Cynoglossoideae are further divided into two and eight tribes, respectively. ...
Article
Pollen, the microgametophyte of seed plants, has an important role in plant reproduction and, therefore, evolution. Pollen is variable in, for example, size, shape, aperture number; these features are particularly diverse in some plant taxa and can be diagnostic. In one family, Boraginaceae, the range of pollen diversity suggests the potential utility of this family as a model for integrative studies of pollen development, evolution and molecular biology. In the present study, a comprehensive survey of the diversity and evolution of pollen from 538 species belonging to 72 genera was made using data from the literature and additional scanning electron microscopy examination. Shifts in diversification rates and the evolution of various quantitative characters were detected, and the results revealed remarkable differences in size, shape and number of apertures. The pollen of one subfamily, Boraginoideae, is larger than that in Cynoglossoideae. The diversity of pollen shapes and aperture numbers in one tribe, Lithospermeae, is greater than that in the other tribes. Ancestral pollen for the family was resolved as small, prolate grains that bear three apertures and are iso‐aperturate. Of all the tribes, the greatest number of changes in pollen size and aperture number were observed in Lithospermeae and Boragineae, and the number of apertures was found to be stable throughout all tribes of Cynoglossoideae. In addition, the present study showed that diversification of Boraginaceae cannot be assigned to a single factor, such as pollen size, and the increased rate of diversification for species‐rich groups (e.g. Cynoglossum) is not correlated with pollen size or shape evolution. The palynological data and patterns of character evolution presented in the study provide better resolution of the roles of geographical and ecological factors in the diversity and evolution of pollen grains of Boraginaceae, and provide suggestions for future palynological research across the family.
... Boraginales are the only order that has more than one family with stinging species. It is composed of 11 families [53], from which three families, i.e., Hydrophyllaceae, Heliotropiaceae and Namaceae, present species with stinging trichomes ( Table 2). Hydrophyllaceae have 12 genera, of which only Phacelia, the largest and most diverse genus of the family (ca. ...
... digynum, H. strigosum and H. subulatum) of the four species analyzed have stinging trichomes [58]. The other genera Euploca, Ixorhea and Myriopus were described as not having stinging trichomes [49,53]. However, it is possible to observe leaf trichomes similar to the stinging ones in a picture of Myriopus embedded in a study of foliar anatomy, although the authors have concluded that such trichomes were absent [74]. ...
Article
Full-text available
Stinging trichomes are rare in plants, occurring only in angiosperms, where they are reported for a few genera belonging to six families. Although there is no report of stinging trichomes in Apocynaceae, previous fieldwork collections of Fischeria and Matelea caused us a mild allergic reaction on the skin when we contacted the dense indumentum of the plants. This fact associated with the well-known presence of glandular trichomes with acute apex in both genera raised suspicions that stinging trichomes could be present in the family. Hence, this study aimed to investigate the likely occurrence of stinging trichomes in Fischeria and Matelea. We analyzed vegetative shoots and leaves of Fischeria stellata and Matelea denticulata through the usual procedures of light and scanning electron microscopy. We also performed several histochemical tests to investigate the chemical composition of trichome secretion. We detected that glandular trichomes occur throughout the surface of the leaf and stem. They are multicellular, uniseriate with an apical secretory cell, which has a dilated base and a needle-shaped apex. The secretion is compressed into the acuminate portion of the apical cell by a large vacuole, and crystals are deposited in the cell wall in a subapical position, providing a preferential site of rupture. The secretion, composed of amino acids and/or proteins, is released under mechanical action, causing skin irritation. Based on our detailed morphological and anatomical analyses, and in the functional aspects observed, we concluded that the glandular trichomes in Fischeria and Matelea can indeed be classified as stinging. Thus, Apocynaceae is the seventh family for which this type of trichome has been reported. We also compiled information on stinging trichomes in all families of angiosperms. Their phylogenetic distribution indicates that they have evolved at least 12 times during angiosperm evolution and may represent an evolutionary convergence of plant defense against herbivory.
... флоры Узбекистана. Авторы придерживаются понимания объема семейств в рамках современной системы Boraginales (Chacon et al., 2016;Luebert et al., 2016). ...
Article
Full-text available
Исследования проведены в рамках проекта «Флора Узбекистана», который представляет собой переход от классического этапа существования ботанической информации, доступной в виде гербарных коллекций и обобщающих их бумажных публикаций, к современному этапу электронного накопления, обработки и распространения информации. При осуществлении критической ревизии видов семейств Heliotropiaceae и Boraginaceae для очередного тома «Флоры Узбекистана» важным этапом работы служит выделение типовых образцов и проведение типификации. Cтатья содержит сведения о 100 типовых образцах названий 38 видов из семейств Heliotropiaceae и Boraginaceae, хранящихся в коллекции Среднеазиатского отдела Национального гербария Узбекистана Института ботаники АН Республики Узбекистан (TASH), 22 образцах, хранящихся в Гербарии Ботанического института им. В. Л. Комарова РАН (LE), 6 образцах из Гербария им. Д. П. Сырейщикова биологического факультета МГУ (MW), и 2 образцах, хранящихся в коллекции Muséum National d’Histoire Naturelle, Paris, France (P). Для каждого типового образца приведены категория, цитата оригинальной этикетки, данные протолога, принятое правильное название и необходимые примечания. Всего указываются 130 типовых образцов, из них 25 лектотипов, 29 изолектотипов, 35 синтипов, 12 голотипов, 7 изотипов, 6 паратипов и 17 автентиков. Обозначены лектотипы названий 13 видов из родов Cynoglossum, Lappula, Macrotomia, Nonea, Onosma, Paracaryum, Rindera, Stephanocaryum. Для 7 таксонов подтвержден выбор гербарного образца в качестве лектотипа. В коллекции типовых образцов представлены виды, которые описали М. Г. Попов, К. З. Закиров, В. К. Пазий, А. П. Чукавина, Е. Г. Черняковская, Р. В. Камелин, А. Д. Ли с территории всех республик Средней Азии: Казахстана, Узбекистана, Таджикистана, Кыргызстана и Туркменистана. Самые ранние сборы в типовой коллекции принадлежат Г. С. Карелину и И. П. Кирилову
... Cordiaceae comprises ca. 400 species distributed in two genera: Cordia L. and Varronia P. Br. (Luebert et al., 2016). Cordia is the largest genus, with ca. ...
Article
Varronia polycephala is a medicinal plant used as a substitute of V. curassavica (the popular “erva baleeira”) in some traditional communities. This paper aims to describe and illustrate the leaf anatomy of V. polycephala. The results show that V. polycephala presents a distinct type of non-glandular trichome in abundance on the abaxial leaf surface, as well as a single type of glandular trichome on the same surface. Other characteristics observed in V. polycephala include non-glandular trichomes with cystoliths, crystal sand idioblasts, shape of the petiole in cross section, type of vascular bundles and ring collenchyma in the petiole. Histochemical results suggest similarities with V. curassavica, showing the pharmacological potential of V. polycephala. Possible ecological functions of structures and substances are discussed.
... The genus Johnstonella Brand (Boraginaceae s. str., after Chacón et al. 2016 andLuebert et al. 2016) was originally segregated from Cryptantha Lehmann ex G.Don and described with two species: Johnstonella inaequata (I.M.Johnst.) Brand and J. racemosa (A.Gray) Brand, the latter the lectotype of the genus (Simpson et al. 2014). The genus was not accepted by subsequent botanists, however, until the molecular phylogenetic study by Hasenstab-Lehman and Simpson (2012). ...
Article
Full-text available
Citation: Simpson MG, Hasenstab-Lehman K, Mabry ME, Muñoz-Schick M (2022) Johnstonella punensis (Boraginaceae), a new species endemic to the dry Puna of Chile. PhytoKeys Abstract In an earlier molecular phylogenetic study, a sample of what was originally identified as Cryptantha hispida (Boraginaceae) from Chile, grouped with species of the genus Johnstonella. This sample was subsequently shown not to be C. hispida, but an undescribed species, endemic to the dry Puna of Chile. This new species is described here as Johnstonella punensis, along with a key to all South American species of the genus. Johnstonella punensis resembles other members of that genus in having an ovate fruit shape, ovate nutlets and a long style that extends beyond the nutlets. It is unusual in the genus in having a non-tuberculate, dimpled to rugulose nutlet surface sculpturing. Its closest relative within the genus is likely the South American J. diplotricha. Resumen En un estudio filogenético molecular anterior, una muestra que originalmente se identificó como Cryptantha hispida (Boraginaceae) de Chile se agrupaba con especies del género Johnstonella. Posteriormente se demostró que esta muestra no era C. hispida sino una especie no descrita, endémica de la Puna seca de Chile. Esta nueva especie se describe aquí como Johnstonella punensis, junto con una clave para todas las especies sudamericanas del género. Johnstonella punensis se parece a otros miembros del género por tener un fruto de forma ovadа, clusas ovadas y un estilo largo que sobrepasa las clusas. Es inusual en el género que la clusa tenga una superficie no tuberculada, sino que rugulosa formando hoyuelos. Su pariente más cercano dentro del género es probablemente la especie sudamericana J. diplotricha. A peer-reviewed open-access journal Michael G. Simpson et al. / PhytoKeys 197: 149-164 (2022) 150
... The taxonomic circumscription of families follows PPG I (2016) for ferns and fern allies, Christenhusz et al. (2011b) for Gymnosperms, and Angiosperm Phylogeny Group (2016) for Angiosperms, with the exception of Dipsacales (Reveal 2011), Caryophyllales (Hernández-Ledesma et al. 2015 and Boraginales (Luebert et al. 2016). Authors' citations of plant names were standardized following the Rec. ...
Article
Full-text available
Digital data concerning the flora of Italy are largely fragmented among different resources hosted on different platforms, and often with different data standards, which are neither con-nected by a common access point, nor by web services, thus constituting a relevant obstacle to data access and usage. Taxonomic incongruences add a further complication. This paper de-scribes Floritaly, an online information system which allows to access and query updated in-formation on the checklist of the flora of Italy, aiming at becoming an aggregator for Italian botanical resources. Floritaly was developed in a collaborative effort by more than 50 taxon-omists, with the support of the Italian Botanical Society, and of Project Drydes (University of Trieste), to provide a better and reliable organization of botanical knowledge in Italy, as well as a relevant simplification for data retrieval, and a further stimulus towards a more collabora-tive approach in botanical research.
... Gottschling & al. (2014) showed that Lennoa and Pholisma were a component of a monophyletic Ehretiaceae, a split-off from Boraginaceae. More recently, Luebert & al. (2016) used additional molecular phylogenetic data as well as morphological features to propose a consensus classification where the order consists of eleven monophyletic families. Here Lennoaceae is wellsupported as sister to Ehretiaceae. ...
Article
Full-text available
Angiosperms that morphologically and physiologically attach to other flowering plants by means of a haustorium have evolved 12 times independently resulting in 292 genera and ca. 4750 species. Although hemiparasites predominate, holoparasitism has evolved in all but two clades, Cassytha (Lauraceae) and Krameria (Krameriaceae). Santalales contains the largest number of genera (179) and species (2428) among the 12 parasitic plant lineages whereas Orobanchaceae is the largest single family with 102 genera and over 2100 species. This review presents the current state of knowledge on the molecular phylogenetic relationships among all clades of parasitic angiosperms. These methods have been particularly important in revealing the closest non‐parasitic relatives of holoparasites, plants that exhibit reduced morphologies, increased substitution rates, and frequent horizontal gene transfers, all of which confound phylogenetics. Although comprehensive molecular phylogenies are still lacking for many of the large genera, nearly complete generic level sampling exists, thus allowing unprecedented understanding of the evolutionary relationships within and among these fascinating plants.
... Boraginaceae s.s. (sensu Luebert et al., 2016;Weigend et al., 2016) are the largest family of Boraginales. Boraginaceae, as used here, are considered the family corresponding to Boraginoideae sensu APG IV (2016). ...
Article
Morphological studies rarely address floral organ modifications or their integration into floral architecture and floral function. Boraginaceae show two prominent types of stamen–corolla tube modifications: faucal and basal scales. Both types, especially faucal scales, are widely used in classification. Here, the ontogeny and morphology of faucal and basal scales are studied in 29 species from eight tribes and all three subfamilies of Boraginaceae s.s. (=Boraginoideae sensu APG IV) using scanning electron microscopy. Integration into floral architecture is visualized with micro-computed tomography (µCT). Faucal and basal scales are present in 18 and 27 species, respectively. Both types of scales develop late in flower ontogeny, but with variable timing. Faucal scales are morphologically far more variable than basal scales. Faucal scales are located close to the anthers and sometimes are involved in anther cone formation. Basal scales cover either the gynoecial disc nectary or the entire ovary. The different scale morphologies identified here enclose complex internal spaces, but they show no obvious phylogenetic patterns. This probably indicates strong functional constraints and adaptive pressures. This is a first in-depth study of the morphology and development of stamen–corolla tube modifications in Boraginaceae s.s., demonstrating that in situ three-dimensional visualization of floral architecture with µCT provides unprecedented insights into the evolution and functional integration of stamen–corolla tube modifications in Boraginales.
... The family Boraginaceae s.l. (e.g., as in APG IV 2016) has been split into several, separate families (see Luebert et al. 2016), including a reduced Boraginaceae s.s., this classification accepted here-henceforth cited as simply Boraginaceae. Chacón et al. (2016) proposed an infrafamilial classification of the Boraginaceae, dividing it into three subfamilies, ten tribes, and six subtribes. ...
Article
Full-text available
Based on a previous molecular phylogenetic analysis, Cryptantha, an herbaceous plant genus of the family Boraginaceae, subtribe Amsinckiinae, was split into five genera: Eremocarya, Greeneocharis, Johnstonella, Oreocarya, and a reduced Cryptantha, the last in two separate clades. As a result of this study, Johnstonella was expanded to 13 species and 15 minimum-rank taxa, these formerly classified in Cryptantha s.l. More recent analyses of this complex, with an increased sample size and high-throughput sequence data, indicate that four additional Cryptantha species not previously sampled-C. albida, C. mexicana, C. texana-plus what was originally identified as C. hispida nest within Johnstonella with strong support. However, the identity of C. hispida used in this analysis is now in doubt. The material used likely represents a new species, in the process of being investigated. Two additional species not sequenced to date-C. geohintonii and C. gypsites-are clearly close relatives of C. albida and C. mexicana, based on morphological similarity. In order to maintain monophyly of genera, we here make new combinations in transferring four of these species from Cryptantha to Johnstonella, with the new combinations Johnstonella albida, J. geohintonii, J. gypsites, and J. mexicana. We delay the transfer of Cryptantha texana to Johnstonella because of its morphological similarity to other species that clearly nest within Cryptantha s.s. These same molecular phylogenetic studies may also support the transfer of two previously recognized Johnstonella species-J. echinosepala and J. micromeres-to Cryptantha, one to each of two separate clades. Additional phylogenetic studies focusing on some of these taxa are needed to confirm the position of these latter three species and the possible recognition of a new genus in the complex.
... The angiosperm list follows the APG IV (2016) classification system, with exception of Cordiaceae and Heliotropiaceae kept apart from Boraginaceae s.l. (BWG 2016) and Passifloraceae s.s. For ferns, Smith et al. (2006) and Rothfels et al. (2012) were followed, while for the lycophytes, Kramer & Green (1990). ...
Article
Full-text available
This study presents a floristic survey of the vascular plants of restingas of Lençóis Maranheses National Park, in the municipality of Barreirinhas, Maranhão, Brazil, including descriptions of the principal phytophysiognomies and similarity analyses including other restinga areas in North and Northeastern Brazil. Samples from the study area deposited in the herbaria MG, IAN and MAR were inventoried and fieldwork for the collection of additional botanical samples was undertaken between September 2015 and August 2017. Unweighted pair group method with arithmetic mean (UPGMA) using Jaccard index was used to calculate the similarity among floras of the restingas of the states of Pará, Piauí, Ceará and Maranhão. A total of 289 species in 189 genera and 73 families was recorded, including 56 new occurence records for the state of Maranhão. The richest families were Poaceae, Cyperaceae, Fabaceae, Rubiaceae, Eriocaulaceae and Myrtaceae. Similarity indexes revealed low similarity among the selected areas, suggesting high floristic identity for each area, as well as possible collection bias among the areas.
... The bulk of small or monotypic genera previously considered to be distinct in the subtribe, viz. Cynoglossopsis Brand, Ivanjohnstonia Kazmi, Paracynoglossum Popov, and Pardoglossum Barbier & Mathez were included in the genus Cynoglossum in Weigend et al. (2016) and Luebert et al. (2016). However, the phylogenetic relationships of the other segregates Paracaryum, Mattiastrum, Rindera, Trachelanthus, Lindelofia and Solenanthus remain to be investigated. ...
Article
Full-text available
Cynoglossinae with ca. 200 spp. is one of the taxonomically most challenging subtribes of tribe Cynoglosseae with regards to generic delimitations. Here, we present a phylogenetic analysis based on nuclear ITS and cpDNA trnL-F and rps16 sequences of 270 accessions including 102 newly sequenced ones, representing all currently recognized genera of Cynoglossinae to clarify relationships among Cynoglossum and allied genera. We conducted Maximum Likelihood, Maximum Parsimony and Bayesian Inference analyses on separate and combined datasets. Cynoglossinae is recovered as a monophylum of closely related genera. Subtribe Cynoglossinae falls into three major groups: (1) clade I including Microparacaryum spp., (2) clade II comprising two species of Lindelofia, several Cynoglossum and Paracynoglossum spp., and (3) a larger clade (clades III+IV) consisting of Solenanthus, Rindera, Trachelanthus, Paracaryum, Pardoglossum, Mattiastrum and the remainder of Cynoglossum. None of the genera were monophyletic in the combined nuclear-plastid and nuclear datasets, except for Microparacaryum. A close relationship between Solenanthus apenninus and Cynoglossum officinale is shown, which suggest transferring S. apenninus to Cynoglossum. Moreover, Pardoglossum atlanticum, the type of the genus, is placed within Mediterranean Cynoglossum corroborating its synonymy under Cynoglossum. Lindelofia olgae is distantly related to the type of the genus (L. longiflora), but more closely connected with Solenanthus core-group. The majority of species currently recognized as members of Rindera do not form a clade with the type of the genus (R. tetraspis). The genus Paracaryum as currently circumscribed is paraphyletic and includes some species of Mattiastrum and Rindera.
... Recently, Hilger & Diane (2003) rearranged the generic delimitation and classification of Boraginaceae subfam. Heliotropioi deae (by some, e.g., Luebert et al. 2016, recognised at family level), based on molecular data of nuclear ITS1 and chloroplast trnLUAA intron sequences, combined with morphological characters. They proposed to merge Schleidenia Endl. ...
Article
Full-text available
In order to recognise both taxa previously regarded as varieties of Heliotropium baclei, nowadays classified in Euploca, a new combination is necessary. As the two varieties are clearly separable in terms of morphology and biogeography, we propose to raise these varieties to the species level, for which the new combination Euploca katangensis needs to be created. Moreover, we propose the new combination Euploca madagascariensis for Heliotropium madagascariense, a species from Madagascar considered by some as conspecific with H. baclei, but treated here as distinct. For these three species with beaked fruits, constituting the ‘Euploca baclei complex’, a key and a distribution map, based on revised herbarium specimens, is given. Two additional combinations, Euploca bullockii and Euploca sessilistigma are made to complete the transfer of tropical African Heliotropium species that belong in Euploca.
... For subsequent molecular dating analyses, the initial dataset of 70 accessions (Supplementary Fig. S3 1 ) was reduced to 56 accessions (one accession per species). Boraginaceae have an abundant fossil record (Weigend et al. 2010;Luebert et al. 2016), but fossils within Cynoglossinae have not yet been unearthed. Therefore, the divergence time was estimated based on a secondary calibration from a previous study by Chacón et al. (2017). ...
Article
The Irano-Turanian (I-T) bioregion harbours one of the Old world’s greatest repositories of botanical diversity; however, the diversification patterns and the phenotypic evolution of its flora are sorely understudied. The subtribe Cynoglossinae is characteristic of the western I-T bioregion, species–rich both in the desertic lowlands and the more mesic highlands of the Iranian plateau. About 70 species of Cynoglossinae are present in the Iranian plateau, 47 of which are endemic to the plateau.Herein, nuclear ITS and cpDNA rpl32-trnL and trnH–psbA sequences were used to investigate the molecular phylogeny, historical biogeography and ancestral character states of Cynoglossinae. Molecular dating and ancestral range reconstruction analyses indicated that the subtribe Cynoglossinae has initiated its diversification from the eastern part of the western I-T during the mid-Miocene, concomitantly with the uplift of the Pamir and Hindu Kush mountains. Moreover, from the Pliocene onwards the Afghan-India collision and extensive deformation of the Arabia-Eurasia convergence probably promoted allopatric speciation in Cynoglossinae via mostly vicariance events. Evolution of annuals with small nutlets from perennials with large nutlets was accompanied by mesic to desert habitats shifts. Herein, to explain distribution of Cynoglossinae in the western I-T, the congruence between cladogenetic, geological and palaeoclimatic events was investigated.
... We aligned the sequences using MAFFT v.7.310 (Katoh & Standley 2013, 2016 and removed all columns in the alignment with 90% or more gaps using Geneious v.9.1.8. We then inferred a Maximum Likelihood phylogeny using IQ-Tree (Nguyen et al. 2015) via the IQ-Tree server (Trifinopoulos et al. 2016), with automatic model selection using ModelFinder (Kalyaanamoorthy et al. 2017) and 1000 ultrafast bootstrap replicates. ...
Article
Euploca riochiquensis is described and illustrated as a species new to science endemic to the Rio Chico valley in Chuquisaca, Bolivia. Morphologically it resembles E. lagoense in its solitary axillary flowers but differs conspicuously in its shrubby habit. Its phylogenetic position, conservation status and distribution are discussed.
... It should be noted here that this species grows in arid lands [28]. Luebert [33] indicates that the Nonea genus is characterized by usually being annual or perennial plants and rarely trees or shrubs. The calyx of the plant is hairy, the corolla is tubular, and the fruits are nutlet. ...
Article
The present study deals with some morphological and anatomical characteristics of the Nonea echioides(L.) Roem. & Sehult species belonging to Boraginaceae, which is recorded to have spread recently in Kurdistan region of Iraq. This research focused on some of the important morphological characteristics of the stems, leaves, flowers, and fruits and comparing them with other studies of neighboring countries to Iraq. These morphological characteristics were found to be important in isolating the species of the filed. The anatomical features of the epidermis, stomata, and trichomes were also investigated. The study shows that Nonea echioides belongs to C3 plants based on the anatomical features of the leaf. In conclusion, the present study provided means for field identification and taxonomy of the plant.
... Cordia L. is the largest genus of Cordiaceae (Boraginales -following the familial classification proposed in BWG (2016)) comprising around 250 species widely distributed in tropical and subtropical regions, with many species restricted to the Neotropics (Miller & Gottschling, 2007;BWG, 2016). Currently its infrageneric classification comprises six sections: Cordia sect. ...
Article
Full-text available
Cordia weddellii, a species of Cordia sect. Cordia (Cordiaceae, Boraginales) endemic to South America is recorded for the first time in Brazil, growing in Cerrado vegetation in Bahia and Tocantins states. A distribution map, photographs, comments on the habitat and phenology, and a preliminary conservation assessment are provided for the species. Additionally, we provide a lectotype for C. weddellii.
... For example, in recognizing the separate families Amaranthaceae sensu stricto and Chenopodiaceae (which were merged under Amaranthaceae s.l. in all versions of the APG classification) we followed the consensus classification of Caryophyllales accepted by Hernández-Ledesma et al. (2015) and other experts in the group. We also prefer to recognize in Boraginales several separate families (Boraginaceae s.str., Cordiaceae, Ehretiaceae, Heliotropiaceae, Hydrophyllaceae, Lennoaceae, Namaceae, etc.), following Luebert et al. (2016), in contrast to the widely circumscribed Boraginaceae as suggested by APG (see APG IV, 2016, and earlier versions). The portfolio now encompasses the main trilogy of the APP, TPP (Tracheophyte Phylogeny Poster), BPP (Bryophyte Phylogeny Poster), a general overview of the embryophytes (EMB), and about 30 further posters on individual orders and families of angiosperms, cumulatively involving a team of more than 130 botanists. ...
Article
Full-text available
Historically, wallcharts and posters created by botanical illustrators, often highly skilled artists, have played an important role in teaching botany at the university level. Large-scale panels and posters can visualize complex interrelationships and entire stories in a clear and appealing overview in graphs, tables, and diagrams. Carrying this concept of educational tools into the electronic era, the Plant Phylogeny Poster project uses this approach for displaying evolutionary relationships in systematic botany. The Angiosperm Phylogeny Poster (APP) displays, as phylogenetically arranged clades, the orders and families of flowering plants (with orders hyperlinked to APweb, Stevens, 2001-onwards), the Tracheophyte Phylogeny Poster (TPP) families and genera of ferns and gymnosperms, and the Bryophyte Phylogeny Poster (BPP) orders and families of liverworts, mosses, and hornworts. The portfolio currently also includes about 30 posters on individual orders and families of angiosperms. Each group within these evolutionary trees is matched with essentially relevant morphological features, biogeographic occurrences, and other information in compactly condensed text blocks. All posters are freely available online, some in more than 30 languages, coauthored by a team of more than 130 botanists. The posters are regularly updated, current literature is cited. The project is expanding steadily and rapidly.
... Boraginaceae (sensu Weigend et al. 2014;Luebert et al. 2016;Hasenstab-Lehman 2017) are widely distributed, mainly in tropical, subtropical and temperate regions, with a high number of poorly studied taxa in the Americas. These lands are among the most diverse ecoregions in the world and, in line with this, their plant diversity has been a challenge, especially for taxonomically complex and diverse genera such as Cryptantha Lehmann ex G. Don. ...
Article
Full-text available
During the preparation of the treatment of the genus Cryptantha Lehmann ex G. Don for South America, numerous names were identified as needing typification to stabilize their nomenclature. As a result, lectotypes are designated for 11 names and second-step lectotypes for 20 names. Furthermore, supporting information about the type material of the basionyms of four Cryptantha names already typified by Johnston (Eritrichium talquinum Phil., Eritrichium dimorphum Phil., Eritrichium carrizalense Phil., and Eritrichium subamplexicaule Phil.) is provided.
... Varronia is a Neotropical genus with ca. 100 species belonging to the Cordiaceae family (formerly included in Boraginaceae) (Miller and Gottschling, 2007;Weeks et al., 2010, Luebert et al., 2016. Distyly, and rarely subdioecy, are the main sexual systems within the genus and the tubular corolla includes urceolate and campanulate shapes (Opler et al., 1975;Miller and Gottschling, 2007;Stutzman et al., 2012). ...
Article
Heterostyly, the reciprocal position of sexual organs between different morphs within a population, is assumed to promote cross-pollination in hermaphroditic plants. Heterostyly is usually linked to an incompatibility system where only the crosses between the different morphs produce fruit. Although heterostyly has been frequently studied in plant species with specialized pollination systems, few studies have investigated this in species with a broad spectrum of floral visitors. Here, we describe the heterostyly of Varronia spinescens and conducted hand-pollination experiments in the field to determine the incompatibility system. We also analysed perianth morphology, morph-ratio, adaptive inaccuracy in reciprocity and pollinator visitation rate in relation to floral morph and number of open flowers. Our results indicate that V. spinescens is a distylous species with a high degree of reciprocity between the sexual organs, presenting a non isoplethic morph ratio with a bias towards the long style (L-) morph. The incompatibility system is heteromorphic and the studied population received visits by a very high number of visitor and pollinator species. Varronia spinescens is thus the heterostylous species of the genus with the second-highest number of registered pollinators. Our results suggest that in a generalist pollination system with little floral morphological restriction to pollinators, the maintenance of sexual polymorphism is likely associated with a high reciprocity between morphs.
Article
A new combination in Euploca (Heliotropiaceae), E. asperrima, endemic to the Galapagos Archipelago, Ecuador, is proposed herein.
Article
Full-text available
An updated checklist of the genus Moltkia is presented and seven taxa (six species and one subspecies) are recognized from the entire world. In this checklist, the effective place of publication of the accepted names, formal taxonomic status, nomenclature, homotypic and non-homotypic synonyms are identified and verified. Lectotypes are here designated for five names. An identification key, illustrations and a distribution map for Moltkia species are provided.
Article
Three new combinations in Myriopus (Heliotropiaceae) endemics from Brazil are proposed in this paper: Myriopus gardnerianus, M. membranaceus and M. salicifolius.
Article
Full-text available
Two new combinations in Varronia (Cordiaceae, Boraginales) are proposed, both species endemic to Colombia: V. fuertesii and V. ramirezii.
Article
Full-text available
Premise of research. Hydrophyllaceae are characterized by simple floral organization contrasting with a peculiar floral architecture resulting from complex compartments formed by stamen–corolla tube modifications. Additionally, the internal ovary architecture of Hydrophyllaceae shows significant variation, while the developmental trajectory of the gynoecium is relatively conserved. Despite insights from recent studies of the flower and fruit evolution of the family, there are only a few ontogenetic studies of Hydrophyllaceae, and a complete understanding of the underlying processes has not yet been achieved. Methodology. Here, we use scanning electron microscopy and micro–computed tomography to investigate the flower and fruit ontogeny of two genera of Hydrophylleae, Emmenanthe and Pholistoma, with a particular focus on the gynoecium and modifications of the stamen–corolla tube. Pivotal results. Our results complement two previously published data sets, broadening our understanding of Hydrophylleae evolution. Hydrophylleae comprise only a few monotypic or small genera, but their floral evolution appears to be remarkably complex, in terms of both gynoecial structure and perianth modifications. We find 10 stamen–corolla tube modifications, although these may be rudimentary in species previously considered as lacking them altogether (e.g., Emmenanthe). The relative conservation of perianth architecture contrasts with the highly variable internal ovary architecture of Hydrophylleae. There is considerable divergence in ovule/seed number, as well as in the details of (parietal) placentation and septation, and we propose a hypothetical evolutionary series for the internal ovary architectural diversity of Hydrophyllaceae. Conclusions. We propose that—starting from a fairly conserved floral organization—minor heterochronous shifts in both perianth and ovary development can explain most of the morphological diversity found in the flowers and fruits of Hydrophylleae.
Article
Boraginales (the forget‐me‐not order) is a core group within the lamiids clade. However, until now, no genome from Boraginales has been reported, and published transcriptomes are also rare. Here, we reported the first Boraginales species de novo genome (i.e., Echium plantagineum genome) and seven other Boraginales species transcriptomes to probe three issues: 1) Boraginales’ phylogenetic position within the lamiids clade; 2) potential whole genome duplications (WGDs) in Boraginales; and 3) candidate key enzyme genes in the alkannin/shikonin core pathway. The results showed that 1) Boraginales was most likely closer to the Solanales/Gentianales clade than the Lamiales clade, at least based on the single‐copy orthologous genes from genome/transcriptome data; 2) after the gamma (γ) event, Boraginaceae (classified into the Boraginales I clade) probably underwent at least two rounds of WGD, whereas Heliotropiaceae and Ehretiaceae (classified into the Boraginales II clade) likely underwent only one round of WGD; and 3) several candidate key enzyme genes in the alkannin/shikonin core pathway were inferred, e.g., genes corresponding to geranyl cyclase, naphthol hydroxylase and O‐acyl transferase.
Article
Myriopus eulinae (Heliotropiaceae), a new species from northeastern Brazil and associated with the Caatinga, Atlantic Forest, and Caatinga-Cerrado transition phytogeographic domain, is described and illustrated. The species can be morphologically recognized by its discolored leaf blade, lax inflorescences, and strigose to hirsute fruits. We discuss its habitat, flowering and fruiting, taxonomic affinities, distribution, and conservation status.
Poster
Full-text available
HILGER HH, COLE TCH, SELVI F, MELO JIM (2020) PÔSTER DA FILOGENIA DAS BORAGINACEAE (Portuguese version of HILGER/COLE/SELVI: BORAGINACEAE PHYLOGENY POSTER) • classificação inframiliar de Boraginaceae s.str. seguindo as recomendações do Boraginales Working Group • árvore hipotética baseada em dados filogenéticos moleculares (2019); revisões de vários subgrupos em andamento • filogenia geral baseada em Chacón et al. 2016 • nomes supragenéricos após Reveal 2011 ff. e Chacón et al. 2016 • contrária ao APG, mas em conformidade com APweb e outras fontes seminais, nós aqui reconhecemos várias famílias de Boraginales • comprimento dos ramos deliberados, não expressando a atual escala de tempo • os caracteres listados (baseados, em sua maioria, em Kubitzki K et al., FGVP Vol. 14 e Chacón J et al. 2016)
Article
Four new combinations in Euploca from South America (Heliotropiaceae), all them endemic to Peru, are proposed in this paper: Euploca lobbii, Euploca oxyloba, Euploca polyanthella, and Euploca toratensis.
Poster
Full-text available
HILGER HH, COLE TCH, SELVI F, OTERO A (2020) PÓSTER DE LA FILOGENIA DE BORAGINACEAE - Versión en español de: HILGER HH, COLE TCH, SELVI F (2020) Boraginaceae Phylogeny Poster • clasificación infrafamiliar de Boraginaceae s.str. siguiendo las recomendaciones del Grupo de Trabajo de Boraginales • árbol hipotético basado en datos filogenéticos moleculares (2019); revisiones bastantes subgroupos en proceso • el grueso de la filogenia está basada en Chacón et al. 2016 • nombres supragenéricos aceptados después de Reveal 2011 ff. y Chacón et al. 2016 • a diferencia de APG, pero en congruencia con APweb y otras fuentes esenciales, se reconocen aquí varias familias dentro de Boraginales • longitudes de rama deliberadas, no expresan la escala temporal real • los caracteres elegidos (ppalm. de Kubitzki K et al., FGVP Vol. 14 y Chacón J et al. 2016)
Article
Full-text available
Cordia obtusiloba, a new species of Cordia section Gerascanthus (Cordiaceae), currently known to the hypoxerophytic caatinga of the state of Sergipe, Brazil, is described and illustrated. A distribution map, data on phenology, and conservation assessment are provided.
Article
Two new combinations in Euploca (Heliotropiaceae) from Marquesas Islands, endemic from French Polynesia, are proposed in this paper: Euploca marchionica and Euploca perlmanii.
Article
The present study provides a taxonomic synopsis of Cordiaceae and Heliotropiaceae (Boraginales) for Uruguay. In order to gain full comprehension of the Uruguayan species in these groups, we examined collections at MVFA, MVJB and MVM, CTES, and SI, consulted virtual collections at F, K, NL-L, NL-U, and P, and studied the online collections hosted by the Herbrio Virtual da Flora e dos Fungos-Reflora. We accept 19 species, six of which belong to Cordiaceae (Cordia and Varronia, each with three species) and 13 to Heliotropiaceae (Euploca with four species, Heliotropium with seven species, and Myriopus with two species). Euploca filiformis and E. krapovickasii are recorded for the first time for Uruguay. We provide identification keys for the recognition of each family and their species, a list of selected specimens, and comments on the morphology, distribution and phenology of each species.
Article
Boraginaceae s.l sofreu muitas modificações na sua tradicional delimitação que incluía quatro subfamílias Ehretioideae (Mart.) Arnott, Boraginoideae Arnott, Cordioideae (Link.) Cham. e Heliotropioideae (Schrad.) Arnott (Judd et al. 2009). Atualmente, com base em dados moleculares, Boraginaceae s.str. está inserida na ordem Boraginales, junto com Codonaceae, Coldeniaceae, Cordiaceae, Ehretiaceae, Heliotropiaceae, Hoplestigmataceae, Hydrophyllaceae, Lennoaceae, Namaceae e Wellstediaceae (BWG 2016).Na área de estudo ocorrem às famílias Cordiaceae e Heliotropiaceae (BWG, 2016). Cordiaceae possui 400 espécies distribuídas por todo mundo, especialmente na região tropical e subtropical (BWG, 2016). São arbustos, subarbustos ou raramente lianas, com codilédones plicados e, geralmente, estiletes duas vezes dicotômicos com quatro estigmas, características exclusivas da família dentro das Boraginales (BWG 2016). Essa família possui apenas dois gêneros: Varronia P. Brown e Cordia L. (Miller & Gottschling 2007). Em Varronia as inflorescências são em glomérulos ou espigas e Cordia em panículas. No Brasil esses gêneros totalizam 87 espécies, sendo Varronia com 32 e Cordia com 55 espécies (Flora do Brasil 2020, em construção).A família Heliotropiaceae possui 450 espécies com quatro gêneros Euploca Nutt., Heliotropium L., Ixorhea Fenzl, Myriopus Small e Tournefortia L. (Weigend & al., 2014).O gênero Ixorhea é o único que não ocorrer no Brasil, pois é endêmica da Argentina (Diane, 2002). A família é caracterizada principalmente por apresentar inflorescências em monocásios escorpioides e estigma inteiro, ocorrem em todo mundo, especialmente nas regiões tropical e subtropical (BWG 2016). Os gêneros mais representativos no Brasil é Euploca Nutt. com 17 espécies e Heliotropium L. com nove espécies (Flora do Brasil 2020, em construção).Este trabalho tem como objetivo realizar o levantamento taxonômico das espécies de Cordiaceae e Heliotropiaceae na região da Chapada Diamantina, Bahia, Brasil. Na Bahia são registrados para Boraginaceae s.l., sete gêneros e 63 espécies (Flora do Brasil 2020, em construção), sendo os gêneros mais diversos Cordia (22) e Varronia (16). Apesar desta grande diversidade, não existe na Bahia especialista no grupo e são escassos os taxonomistas que trabalham com essas famílias no Nordeste, apenas o Dr. José Iranildo Miranda Melo, na Paraíba. Isso implica no desconhecimento da real diversidade do grupo e justifica a existência de várias coletas sem identificação nos herbários, como por exemplo no HUEFS (Herbário da Universidade Estadual de Feira de Santana) existem 44 exsicatas sem identificação, na rede PPBio apenas quatro espécies estão identificadas o que é um número muito baixo para a diversidade do grupo na região.
Chapter
The eudicots are a large, monophyletic assemblage of angiosperms, comprising roughly 190,000 described species, or 75% of all angiosperms. The monophyly of eudicots is well supported from molecular data and delimited by at least one palynological apomorphy: a tricolpate or tricolpate-derived pollen grain. A tricolpate pollen grain is one that has three apertures, equally spaced and approximately parallel to the polar axis of the grain. Apertures are differentiated regions of the pollen grain wall that may function as the site of pollen tube exitus as well as to allow for expansion and contraction of the pollen grain with changes in humidity. Tricolpate pollen grains evolved from a monosulcate type (having a single distal aperture, which is considered to be ancestral in the angiosperms, as well as for many seed plant clades. Many eudicots have pollen grains with more than three apertures, of a great variety of numbers, shapes, and position (constituting important taxonomic characters). These are all thought to have been derived from a tricolpate type.
Article
Full-text available
As a result of the studies, it was revealed that on the territory of Uzbekistan Heliotropiaceae family is represented by 2 genera and 15 species and Boraginaceae family is represented by 31 genera and 118 species, distributed in 10 tribes. The largest number of species is represented by the genera from the tribes Eritirchieae (31) and Cynoglosseae (25). Tribe Lithospermeae (21) is distinguished by specificity of the species composition. The most ancient tribes Boragineae (8), Echieae (3), Trichdesmeae (3) are represented by a small number of taxa.
Article
A complete nomenclatural synopsis is provided for Onosma (Boraginaceae) and the allied genera Maharanga and Podonosma, covering each name of new taxa, each new combination and each replacement name communicated since 1753 till October 2021. All the 749 validly published names are listed together with an account of the formal reasons why 136 additional names must be regarded as invalid. Type species are indicated or newly designated for all the infrageneric names. Information is assembled on the types of the 466 names of specific and infraspecific taxa retrieved from a comprehensive survey of the scientific literature and from public databases, with 126 names (27%) lectotypified here. Finally, 42 new combinations are proposed to solve as yet unresolved nomenclatural issues.
Article
Full-text available
The Early Cretaceous (145–100 million years ago (Ma)) witnessed the rise of flowering plants (angiosperms), which ultimately lead to profound changes in terrestrial plant communities. However, palaeobotanical evidence shows that the transition to widespread angiosperm-dominated biomes was delayed until the Palaeocene (66–56 Ma). Important aspects of the timing and geographical setting of angiosperm diversification during this period, and the groups involved, remain uncertain. Here we address these aspects by constructing and dating a new and complete family-level phylogeny, which we integrate with 16 million geographic occurrence records for angiosperms on a global scale. We show substantial time lags (mean, 37–56 Myr) between the origin of families (stem age) and the diversification leading to extant species (crown ages) across the entire angiosperm tree of life. In turn, our results show that families with the shortest lags are overrepresented in temperate and arid biomes compared with tropical biomes. Our results imply that the diversification and ecological expansion of extant angiosperms was geographically heterogeneous and occurred long after most of their phylogenetic diversity originated during the Cretaceous Terrestrial Revolution.
Article
Full-text available
PREMISE: Fruit type and morphology are tightly connected with angiosperm diversification. In Boraginales, the first-branching families, including Hydrophyllaceae, have one-to many-seeded capsules, whereas most of the remaining families have four-seeded indehiscent fruits. This fact argues for many-seeded capsules as the ancestral condition. However, little is known about the evolution of fruit dehiscence and seed number. The present study investigated the gynoecium and fruit development and morphology and the evolution of seed-numbers in Hydrophyllaceae. METHODS: Gynoecium and fruit development and morphology were studied using scanning electron microscopy and x-ray microcomputed tomography. Ancestral character state reconstruction of seed number was performed using a broadly sampled phylogeny of Boraginales (ndhF and ITS) with an emphasis on Hydrophyllaceae. RESULTS: Our ontogenetic studies not only demonstrate parallel developmental trajectories across Hydrophyllaceae, but also a striking diversity regarding the internal organization of the gynoecium. Ovule number appears to determine ovary structure. Many-seeded capsules are retrieved as the ancestral state of Hydrophyllaceae. At least seven transitions to fruits with (one to) four seeds and four reversals (i.e., from four-to many-seeded fruits) were reconstructed in Hydrophyllaceae. CONCLUSIONS: Several shifts in seed number from “many” to “four” and back to “many” have taken place in capsular-fruited Hydrophyllaceae, a strikingly high number considering that seed number is virtually conserved across the rest of the order. The groups with a conserved seed number of four are characterized by indehiscent schizocarps or drupes and by seeds that are integrated into mericarps. This functional integration probably acts as an evolutionary constraint to shifts in seed number.
Article
This paper includes studies on Heliotropium marifolium complex from India. Five new combinations in Euploca from H. sect. Orthostachys are proposed here. H. marifolium, H. marifolium subsp. wallichii, H. rottleri, H. cornutum and H. madurense are transferred to the genus Euploca. The taxonomic status of the unresolved name H. rottleri is also discussed.
Poster
Full-text available
HILGER HH, COLE TCH, SELVI F, SONG JH 송준호 (2021) 지치과의 계통수, Korean version of: HILGER, COLE, SELVI (2020) Boraginaceae Phylogeny Poster •지치목 워킹그룹(Boraginales Working Group)의 제안에 따른 협의의 지치과 분류체계 •분자계통학적 자료(2019년 1월)에 기반한 가설 계통수; 몇몇 서브그룹(하위집단)의 분류학적 개정 작업은 현재 진행 중에 있음 • Chacon et al. (2016)에 근거한 전체 계통수 •속보다 상위의 학명은 Reveal (2011)과 Chacon et al. (2016) 이후의 학명을 따름 • APG (Angiosperm Phylogeny Group)와는 상충하지만, APweb (Angiosperm Phylogeny Website)과 그 밖의 중요한 자료에 따라, 우리는 본 계통수에서 지치목 내 몇몇의 과를 인식함 •계통수의 절간 길이는 실제 시간의 척도를 반영하지 않은 인위적인 것임 •제시된 형질[주로 Kubitzki K et al., FGVP Vol. 14, Chacon J et al. (2016)을 참조하여 반영]은 각 분계조의 모든 분류군에 반드시 공통적으로 적용되지 않음
Article
“Dust seeds” with an undifferentiated embryo and unipolar germination are produced by holoparasitic species in 10 families of angiosperms. However, aside from a few Orobanchaceae species important in agriculture (crop weeds) and in Chinese traditional medicine we know little about seed germination in these achlorophyllous plants. Our primary purpose was to review the literature on seed germination in nine families of the mostly economically-nonimportant taxa of holoparasites and (briefly) of Cynomoriaceae, an economically important medicinal family for which several publications on its germination are in Chinese. A second purpose was to pull together information on sizes of dust seeds and their undifferentiated embryos and endosperm in both economically- and noneconomically-important holoparasitic plants. We found that not much progress has been made in understanding dormancy-break/germination in these holoparasites since 1969, when the author of a book on parasitic flowering plants pointed out how little we know about their germination. Lack of progress on germination of dust seeds of these holoparasites is not necessarily due to lack of research but at least in part to their nonresponsiveness to well-known treatments used to break dormancy in seeds of autotrophic and hemiparasitic flowering plants. For most species, seed length is <1.0 mm (in many <0.50 mm) and embryo length < 0.40 mm (in many <0.10 mm); endosperm is scant to copious. We suggest that seed conditioning, an essential step in germinating seeds of weedy Orobanchaceae holoparasites, be added to the protocol for germinating the dust seeds of noneconomically-important holoparasites with an undifferentiated embryo.
Poster
Full-text available
HILGER HH, COLE TCH, SELVI F, MAJESKÝ Ľ, MÁRTONFI P (2020) FYLOGENÉZA ČEĽADE BORAGINACEAE • klasifikácia čeľade Boraginaceae s.str. je podľa odporúčaní „Boraginales Working Group“ • hypotetický evolučný strom založený na molekulárnych fylogenetických dátach (2020); naďalej prebieha revízia niektorých skupín • základné fylogenetické vzťahy vychádzajú z práce Chacón et al. 2016 • mená nad úrovňou rodov vychádzajú z prác Reveal 2011 a nasl. a Chacón et al. 2016 • oproti APG, ale v súlade s APWeb a ďalšími kľúčovými zdrojmi rozlišujeme niekoľko čeľadí v rade Boraginales • dĺžka vetiev je arbitrárna a nevyjadruje skutočnú časovú divergenciu • uvádzané znaky (hlavne z Kubitzki K et al., FGVP Vol. 14 a Chacón J et al. 2016) nemusia byť nevyhnutne prítomné pri všetkých členoch danej skupiny
Article
Full-text available
We describe four new species of Cordia (Cordiaceae) from Brazil: Cordia fusca, C. glabrifolia, C. restingae, and C. tarodae. These are illustrated, and data on habitat, phenology, and distribution are provided.
Article
Full-text available
RESUMO Baseando-se na análise de espécimes advindos de 71 herbários nacionais e estrangeiros, trabalhos de campo e literatura especializada, foram determinados os padrões de distribuição geográfica das espécies dos gêneros Euploca e Heliotropium no Brasil. Foram detectados quatro padrões de distribuição geográfica e onze padrões biogeográficos. O centro de diversidade das espécies de Euploca no Brasil é a Região Nordeste. As espécies do gênero Heliotropium concentram-se, na sua maioria, na Região Sul, com o estado do Rio Grande do Sul representando o centro de diversidade do gênero no país. São apresentadas tabelas, mapas e discussões sobre a distribuição e respectivos padrões biogeográficos reconhecidos.
Article
Full-text available
Premise of the study: Major relationships within Lamiidae, an asterid clade with ∼40000 species, have largely eluded resolution despite two decades of intensive study. The phylogenetic positions of Icacinaceae and other early-diverging lamiid clades (Garryales, Metteniusaceae, and Oncothecaceae) have been particularly problematic, hindering classification and impeding our understanding of early lamiid (and euasterid) character evolution. Methods: To resolve basal lamiid phylogeny, we sequenced 50 plastid genomes using the Illumina sequencing platform and combined these with available asterid plastome sequence data for more comprehensive phylogenetic analyses. Key results: Our analyses resolved basal lamiid relationships with strong support, including the circumscription and phylogenetic position of the enigmatic Icacinaceae. This greatly improved basal lamiid phylogeny offers insight into character evolution and facilitates an updated classification for this clade, which we present here, including phylogenetic definitions for 10 new or converted clade names. We also offer recommendations for applying this classification to the Angiosperm Phylogeny Group (APG) system, including the recognition of a reduced Icacinaceae, an expanded Metteniusaceae, and two orders new to APG: Icacinales (Icacinaceae + Oncothecaceae) and Metteniusales (Metteniusaceae). Conclusions: The lamiids possibly radiated from an ancestry of tropical trees with inconspicuous flowers and large, drupaceous fruits, given that these morphological characters are distributed across a grade of lineages (Icacinaceae, Oncothecaceae, Metteniusaceae) subtending the core lamiid clade (Boraginales, Gentianales, Lamiales, Solanales, Vahlia). Furthermore, the presence of similar morphological features among members of Aquifoliales suggests these characters might be ancestral for the Gentianidae (euasterids) as a whole.
Article
Full-text available
A parsimony analysis of 156 representative sequences of the Asteridae sensu lato and 28 outgroup sequences was conducted using a two-tiered approach. First, an analysis of the entire group, including 105 sequences, examined relationships among major lineages within the Asteridae s.l.; subsequently, several clades within the larger group were examined individually in greater detail by including more sequences for the group in question. The search strategy was designed to discover multiple islands of equal parsimony using the heuristic search routine in PAUP. In the broad search and in each more detailed search of subclades, multiple islands were found that imply substantially different relationships. The results suggest a monophyletic Asteridae s.l., comprising the Ericales, Primulales, Ebenales and relatives of the Dilleniidae sensu Cronquist; Cornales, Apiales, and Hydrangeaceae of the Rosidae sensu Cronquist; and the conventionally circumscribed Asteridae. Within the Asteridae s.l., 11 groups were congruent between islands and are designated as follows: (1) Cornales, (2) ericalean clade, (3) Garrya clade, (4) Ilex clade, (5) Apiales, (6) Dipsacales, (7) Asterales s.l., (8) Gentianales, (9) Solanales, (10) Boraginales, and (11) Lamiales s.l. The only grouping between the level of these 11 clades and the whole Asteridae s.l. that is congruent between islands is the clade consisting of the Gentianales, Solanales, Boraginales, and Lamiales s.l., i.e., the Lamiidae of Takhtajan.
Article
Full-text available
The Lennoaceae are a small family of New World holorhizoparasites distantly related to the Boraginaceae and Verbenaceae. The three genera and total of seven species that have tradi- tionally been accepted for the family are here reduced to two genera and four species. Lennoa is a genus of annual plants with 8-merous flowers, biseriate nodding stamens with basally spreading thecae, 7–12 vascular bundles per stem, pollen exine with a granular infrastructural layer, and diploid with n = 9. Lennoa is monotypic, with two formae of L. madreporoides recognized. Pholisma is a genus of perennial plants with 4–10-merous flowers, uniseriate erect stamens with parallel thecae, 15–25 vascular bundles per stem, tectate-columellate exine infrastructure, and tetraploid with n = 18. Pholisma (including Ammobroma) contains three very well-marked species: P. arenarium, P. culiacanum, and P. sonorae.
Article
Full-text available
Dioecious and thorny Rochefortia Sw. is a poorly known though distinct element of the Ehretiaceae comprising woody plants restricted to the Caribbean and the adjacent American mainland. The approximately ten species display a great morphological variability and overlapping taxonomic boundaries, which makes it difficult to differentiate them (particularly in the Caribbean region). We investigated the phylogenetic relationships of Rochefortia using DNA sequence data from one nuclear locus (Internal Transcribed Spacer) and three chloroplast DNA loci (rps16, trnL–trnF, trnS–trnG). The monophyly of Rochefortia was confirmed, with a sister group relationship between an American mainland clade and a Caribbean clade. The latter segregates into three, morphologically rather variable lineages, distributed either in the Lesser Antilles or in the eastern Greater Antilles or in the western Greater Antilles. Thus, geographic occurrence rather than morphology is indicative of taxonomic delimitation in Rochefortia.
Article
Full-text available
This work points out some variations in leaf anatomy useful in the separation of two species of the genus Varronia P.Br., V. globosa Jacq. and V. leucocephala (Moric.) J. S. Mill., and some anatomical adaptations of the semiarid climate. These species differ in stomata distribution, types of glandular trichomes, non-glandular trichomes density, accumulation of substances in V. leucocephala, crystal types, colenchyma type in midrib and petiole, and the vascular bundles in petiole. As unifying characters, both have uniseriate epidermis, glandular and non-glandular trichomes, dorsiventral leaves, crystals, collateral vascular bundles in leaf blades, and petiole with three vascular traces. The morphological study of trichomes has been extensively explored since it is one of the main characteristics differing the species from the genus, and being recognized several types of glandular trichomes, particular to each species. Some anatomical typical features of plants occurring in xeric environments were also identified: stomatal distribution, abundant trichomes with micropapillae on its surfaces, and lipid accumulation.
Article
Full-text available
A phylogenetic analysis of 589 plastid rbcL gene sequences representing nearly all eudicot families (a total of 308 families; seven photosynthetic and four parasitic families are missing) was performed, and bootstrap re-sampling was used to assess support for clades. Based on these data, the ordinal classification of eudicots is revised following the previous classification of angiosperms by the Angiosperm Phylogeny Group (APG). Putative additional orders are discussed (e.g. Dilleniales, Escalloniales, Vitales), and several additional families are assigned to orders for future updates of the APG classification. The use of rbcL alone in such a large matrix was found to be practical in discovering and providing bootstrap support for most orders. Combination of these data with other matrices for the rest of the angiosperms should provide the framework for a complete phylogeny to be used in macro-evolutionary studies.
Article
Full-text available
O presente trabalho trata do estudo taxonômico do gênero Euploca (Heliotropiaceae) no Brasil, sendo registradas 17 espécies. É apresentada chave para o reconhecimento, descrições, ilustrações e comentários, além de dados de distribuição, habitat, floração e frutificação para as espécies.This work deals with a taxonomic study of the genus Euploca (Heliotropiaceae) in Brazil; seventeen species are recorded. A keyfor identification, descriptions, illustrations and comments, besides distribution, habitat, flowering and fruiting data for the species are presented.
Article
Full-text available
The Lamiidae, a clade composed of approximately 15% of all flowering plants, consists of five orders: Boraginales, Gentianales, Garryales, Lamiales, and Solanales; and four families unplaced in an order: Icacinaceae, Metteniusiaceae, Oncothecaceae, and Vahliaceae. Our understanding of the phylogenetic relationships of Lamiidae has improved significantly in recent years, however, relationships among the orders and unplaced families of the clade remain partly unresolved. Here, we present a phylogenetic analysis of the Lamiidae based on an expanded sampling, including all families together, for the first time, in a single phylogenetic analyses. Phylogenetic analyses were conducted using maximum parsimony, maximum likelihood, and Bayesian approaches. Analyses included nine plastid regions (atpB, matK, ndhF, psbBTNH, rbcL, rps4, rps16, trnL-F, and trnV-atpE) and the mitochondrial rps3 region, and 129 samples representing all orders and unplaced families of Lamiidae. Maximum Likelihood (ML) and Bayesian trees provide good support for Boraginales sister to Lamiales, with successive outgroups (Solanales + Vahlia) and Gentianales, together comprising the core Lamiidae. Early branching patterns are less well supported, with Garryales only poorly supported as sister to the above 'core' and a weakly supported clade composed of Icacinaceae, Metteniusaceae, and Oncothecaceae sister to all other Lamiidae. Our phylogeny of Lamiidae reveals increased resolution and support for internal relationships that have remained elusive. Within Lamiales, greater resolution also is obtained, but some family interrelationships remain a challenge.
Article
Full-text available
Typifications are proposed to facilitate ongoing studies in Cordiaceae and to maintain current usage. Lectotypes are designated for the following 13 names: Cordia ecalyculata, C. digyna, C. diospyrifolia, C. exaltata var. melanoneura, C. magnoliifolia, C. obscura var. tomentosa, C. obscura var. magnifolia, C. salicifolia, C. sellowiana, C. silvestris, C. ulei, C. ucayaliensis, and Patagonula glaziovii. Two new synonyms are proposed: C. crenatifolia Rizzini to C. glazioviana (Taub.) Gottschling & J. S. Mill. and C. araripensis Rizzini to C. bicolor A. DC. Additionally, nomenclatural notes on C. blanchetii are provided.
Article
The plate designated as lectotype of the name Heliotropium arborescens L. (1759) does not permit a precise application of the name. The herbarium material associated with that illustration was examined in order to clarify the identity of the type material and an epitype was selected. The epitype corresponds in morphology to the taxon called Heliotropium urbanianum K. Krause (1906) in the recent literature, which is here lectotypified and synonymized with H. arborescens. The name Heliotropium arborescens has been misapplied to a predominantly Peruvian species, which should now be correctly called Heliotropium corymbosum Ruiz & Pav. (1799). The epitypification here proposed will ensure nomenclatural stability for most material from cultivation, where the name Heliotropium arborescens is widely used.
Article
http://digitalcommons.library.umaine.edu/fac_monographs/1172/thumbnail.jpg
Article
A taxonomic revision is provided for Cordia section Gerascanthus and 23 species are recognized. They are characterized by flowers with tubular, ribbed calyces, marcescent corollas that persist and surround the developing fruits, aiding in their wind dispersal, and single-seeded ellipsoid fruits with a fibrous wall. All of the species are restricted to the Neotropics. Thirteen occur in Mexico, and the remainder are scattered throughout Central America, the West Indies, and South America; however, there is no secondary center of diversity. Classification of the section and its morphology are reviewed, and a key and descriptions are provided for all species along with notes on their distributions and conservation status.
Article
This study revises the taxonomy of Varronia P. Br. (Cordiaceae) species endemic to the Galapagos Islands and presents a key that can more reliably distinguish the species using vegetative characters alone. We conclude that there are four endemic species of Galapagos Varronia distributed across ten islands: V. canescens Andersson (syn = V. anderssonii, nom. illeg,), V. lucophlyctis (Hook, f.) Andersson, V. scouleri (Hook, f.) Andersson, and V revoluta (Hook, f.) Andersson. Although most species share island distributions, species are not found in close proximity together and no obviously intermediate or putatively hybrid individuals were observed. The endemic species appear to be distylous, and preliminary data suggest these species may be obligate outcrossers. The findings of our study will assist efforts by park managers and other researchers to determine the conservation status of each species and to develop appropriate conservation measures in line with the life-history of the species.
Article
The phylogenetic relationships of Cordiaceae were investigated using ITS1 sequences from 50 ingroup taxa and three outgroup taxa. Molecular analyses suggest that Coldenia procumbens, traditionally placed in Ehretiaceae, is more closely allied to Cordiaceae and is sister to their core representatives. Cordiaceae segregate into four major monophyletic assemblages, more or less corresponding to established taxonomic units as previously inferred from morphology, ecology, and biogeography. (1) Varronia (currently recognized as Cordia sect. Varronia) is a well-differentiated taxon and has a sister-group relationship to the other three clades (Cordia s. str.). The (2) Collococcus clade comprises the New World species of Cordia sect. Myxa (excluding Cordia dentata and Cordia lutea), all of Cordia sect. Superbiflorae, and Cordia varronifolia (from Cordia sect. Rhabdocalyx). The (3) Myxa clade includes all of the Old World species of Cordia sect. Myxa and the New World species Cordia dentata and Cordia lutea. The (4) Sebestena clade includes Cordia sects. Cordia and Gerascanthus, three species of Cordia sect. Rhabdocalyx, and Auxemma, Patagonula, and Saccellium. Pollen morphology and wood anatomy lend additional support for the close relationship of Auxemma, Patagonula, and Saccellium with Cordia sebestena and its allies. Thus, Auxemma, Patagonula, and Saccellium are no longer tenable as separate taxa and should be reduced under Cordia. A set of ancestral characters (including apomorphies and plesiomorphies) is reconstructed for Cordiaceae based on the phylogenetic hypotheses as inferred from the congruence between morphological and ITS1 data.
Article
The new species Wellstedia robusta, from mountain ridges and slopes south-west of Qandala in north-eastern Somalia, is described and illustrated.
Article
Lectotypes are designated here for 14 names proposed by Ruiz and Pavón in "Flora peruviana, et chilensis" (1799) that were either described or are currently recognized as members of the genera Heliotropium or Tournefortia (Heliotropiaceae): Heliotropium corymbosum, H. incanum, H. lanceolatum, H. microcalyx, H. microstachyum, H. oppositifolium, H. pilosum, H. synzystachyum, Lithospermum aggregatum, Tournefortia angustiflora, T. longifolia, T. polystachya, T. undulata, T. virgata. Currently accepted names and comments on typifications and taxonomic affinities are also provided.
Article
Boraginaceae have erroneously been associated with helicoid cymes, when upon closer inspection they exclusively possess scorpioid cymes (cincinni). A discussion of the inflorescence structure in Boraginaceae is presented with the help of an overview of monochasial branching systems and a three-dimensional representation thereof. Reasons for the incorrect description of Boraginaceae inflorescences are also provided.
Article
A phylogenetic analysis of a combined data set for 560 angiosperms and seven outgroups based on three genes, 18S rDNA (1855 bp), rbcL (1428 bp), and atpB (1450 bp) representing a total of 4733 bp is presented. Parsimony analysis was expedited by use of a new computer program, the RATCHET. Parsimony jackknifing was performed to assess the support of clades. The combination of three data sets for numerous species has resulted in the most highly resolved and strongly supported topology yet obtained for angiosperms. In contrast to previous analyses based on single genes, much of the spine of the tree and most of the larger clades receive jackknife support ≥50%. Some of the noneudicots form a grade followed by a strongly supported eudicot clade. The early-branching angiosperms are Amborellaceae, Nymphaeaceae, and a clade of Austrobaileyaceae, Illiciaceae, and SchiÍsandraceae. The remaining noneudicots, except Ceratophyllaceae, form a weakly supported core eumagnoliid clade comprising six well-supported subclades: Chloranthaceae, monocots, Winteraceae/Canellaceae, Piperales, Laurales, and Magnoliales. Ceratophyllaceae are sister to the eudicots. Within the well-supported eudicot clade, the early-diverging eudicots (e.g. Proteales, Ranunculales, Trochodendraceae, Sabiaceae) form a grade, followed by the core eudicots, the monophyly of which is also strongly supported. The core eudicots comprise six well-supported subclades: (1) Berberidopsidaceae/Aextoxicaceae; (2) Myrothamnaceae/Gunneraceae; (3) Saxifragales, which are the sister to Vitaceae (including Leea) plus a strongly supported eurosid clade; (4) Santalales; (5) Caryophyllales, to which Dilleniaceae are sister; and (6) an asterid clade. The relationships among these six subclades of core eudicots do not receive strong support. This large data set has also helped place a number of enigmatic angiosperm families, including Podostemaceae, Aphloiaceae, and Ixerbaceae. This analysis further illustrates the tractability of large data sets and supports a recent, phylogenetically based, ordinal-level reclassification of the angiosperms based largely, but not exclusively, on molecular (DNA sequence) data.
Article
A study on four species of the genus Phacelia (Hydrophyllaceae) was focused on flower and fruit development. A combination of morphological characters of the corolla and the gynoecium (including formation and distribution of crystals) is shown to be useful in segregating even closely related taxa within the genus. Half inferior ovaries, as demonstrated for two species, seem to be more frequent within the genus than previously assumed. Corresponding stages during early gynoecium development and characters of the nectary disc and corolla may underline a closer relationship between Hydrophyllaceae and Boraginaceae.
Article
An analysis of evolutionary relationships within the angiosperm family Hydrophyllaceae was completed. A parsimony search using DNA sequence data from the chloroplast gene ndhF from 65 accessions yielded 3,227 most parsimonious trees distributed over six tree islands. Hydrophyllaceae, excluding Hydrolea and Codon, are nested within a paraphyletic Boraginaceae sensu lato. These "core" Hydrophyllaceae are restricted to the New World and have the derived feature of capsular fruits with parietal placentation and many small seeds. Two large clades can be recognized-a clade of annual and perennial herbs including Phacelia, tribe Hydrophylleae, and related genera, and a clade containing Nama and the woody, shrubby genera Eriodictyon, Turricula, and Wigandia. The second clade corresponds to the tribe Nameae sensu Gray, but the first has not been formally recognized. Evolutionary relationships based on ndhF are compared to current classification systems and inferences about evolution based on published morphological and anatomical studies.
Article
Codonaceae (Boraginales) is here proposed as a novel segregate family from Boraginaceae, based on Boraginaceae subfamily Codonoideae. The family consists of a single genus, Codon (2 species, Southwest Africa) which is traditionally considered as the only African representative of the predominantly New World family Hydrophyllaceae. Recent molecular studies clearly indicated that it is more closely allied to Boraginaceae s. str. ("herbaceous" Boraginaceae), than to Hydrophyllaceae, however, Codon is morphologically highly aberrant for both families with its polymerous perianth and androecium, style inserted on the apex of an ovoidal ovary, many-seeded, sub-bilocular, loculicidal capsules, endospermous seeds with a very irregularly reticulate testa, and peculiar spines with a multicellular pedestal and unicellular apex. The genus was therefore recently placed into a monotypic subfamily Codonoideae in a broadly defined Boraginaceae s.l. (i.e., including Heliotropioideae, Hydrophylloideae, Cordioideae as subfamilies). We advocate the recognition of the morphologically well-differentiated clades of Boraginales at family rather than subfamily level and therefore propose the recognition of Codonaceae as a novel segregate family. Boraginales then consists of a total of seven families: Boraginaceae s. str., Codonaceae, Cordiaceae, Heliotropiaceae, Hydrophyllaceae, Lennoaceae and Wellstediaceae.
Article
This project sampled throughout Phacelia using the internal transcribed spacer region (ITS-1, ITS-2, and 5.8S gene) of nuclear ribosomal DNA (nrITS) and the chloroplast DNA gene (ndhF) to infer phylogenies for nuclear and plastid partitions. Nuclear and plastid partitions were incongruent in our analyses. Phylogenetic analyses (maximum parsimony, maximum likelihood, and Bayesian inference) recovered gene tree topologies similar to previous molecular studies. We corroborate incongruence between nuclear and plastid topologies for placement of some problematic groups (e.g., Draperia, Romanzoffia and “core” Phacelia subg. Pulchellae, Phacelia sect. Baretiana). Combined analyses resulted in better resolution than separate analyses, and in a topology that favored the separate plastid topologies. Romanzoffia was sister to a monophyletic Phacelia in the combined analyses. Our results support combining incongruent partitions in a combined analysis to seek support for internal nodes. Maximum likelihood analyses were used to infer ancestral chromosome numbers and identify gains, losses, polyploid doubling, and whole genome duplication events from published chromosome counts in the genus. The predicted base number for the genus was x = 9, x = 11, or x = 12.
Article
The development of flowers and fruits of 23 species of the Cynoglosseae and Eritrichieae, formerly for the most part included in the genus “Echinospermum”, was studied morphologically and histologically by means of SEM and microtome sectioning. 1. The early floral ontogeny is strongly uniform in all species (as well as in two species of the Lithospermeae investigated in addition). The sepal, petal, and stamen primordia display a regular 2/5 spiral phyllotaxis. The two carpels develop very late, when the stamina already exhibit the thecae. Soon the flanks of the carpel primordia fuse and give rise to a gynoecial cone. 2. The four mericarps arise as lateral bulges of the carpels. Thereby the gynoecial cone splits up into a terminal style, an intermediate mericarp-bearing columella, and a basal disk. The latter will apparently act as nectary. The part of the fruit that remains on the plant when the nutlets are released is termed gynobase. 3. Just before the stylar slit closes, a low transverse ridge divides the basal cavity of the gynoecium into two shallow grooves. This ridge is interpreted as the septum. The four ovules develop within the angle between the septum and carpel wall, but are clearly located on the septum. Therefore the placentation is (marginal-)axile. In later developmental stages the funiculi remain united with the carpel walls by intercalary growth, feigning a parietal placentation. 4. From the very first the ovules grow diagonally upwards into the mericarpial cavities. Inside the gynoecium four apical and basal septa narrow the connections between ovarial and mericarpial chambers, leaving only a small tube for each funicle. The abscission layers develop inside these massive diaphragms. 5. False septa, often considered characteristic in the Boraginaceae, do not develop in all species. If present, they originate from subepidermal ventral meristems of the carpels. They are not necessary for the formation of the mericarps. 6. The initial globose shape of the Cynoglosseae nutlets remains until ripening. After bulging up from the gynoecial cone the gynobase grows in horizontal direction. So the nutlets attain a position with their disks parallel with the basal area of the gynoecium. The style seems to insert on the receptacle. A subsequent growth in vertical direction brings the mericarps to their final oblique position. In contrast, in the Eritrichieae a forced vertical growth of the gynobase produces the typical triangular shape and the continuously oblique position of the mericarps. Asperugo and Sclerocaryopsis confirm their isolated position within the Eritrichieae by unusual nutlet arrangement. Neighboring mericarps of different carpels turn to each other. 7. Most members of the Cynoglosseae and Eritrichieae exhibit a thickened rim or broad wing around the disk of the nutlets and/or glochids on the nutlet surface. It may be of systematic importance whether the wings are developed prior to glochids or vice versa. In the latter condition a subsequent peripheral growth of the nutlet margins may produce a broad glochidless rim analogous to a wing. After ontogenetic investigation, genera with similarly looking fruits can be distinguished. Heterocarpy as well as heteromericarpy displaying pairs with different wing and glochid equipment seem to have arisen independently in various genera. 8. A broadened definition for the terms “eremocarps, mericarps, and fruits (Klausen, Klausenfrüchte)” is proposed. It may also be applied to the fruits of some other taxa in which parts of the fruit enclosing seeds break away and leave a gynobase.
Article
Tiquilia is very different from the other members of the Ehretiaceae (Boraginales) in many aspects of morphology and ecology. Because detailed knowledge about flower and fruit traits is necessary to reliably infer character evolution of and within Tiquilia, we investigated flower to fruit ontogeny in eight species of Tiquilia using light and electron microscopy. Tiquilia accumulated a number of autapomorphies such as the prostrate growth form, the lack of lateral and ventral bundles in the gynoecium, and the formation of nutlet-like mericarpids as dispersal units instead of more or less succulent drupes. The internal architecture of the superior bicarpellate ovary resulted from the development of several secondary septa including apical, basal and false septa, as it has been reported also from other Boraginales. However, no character found in Tiquilia can be regarded as synapomorphic with any other taxon of the Ehretiaceae. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, ●●, ●●–●●.