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Familial classification of the Boraginales
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.
... Boraginales are a group of flowering plants consisting of up to 2700 species of trees, shrubs, or (often) herbs, with a nearly worldwide distribution (Mabberley, 2008;Luebert & al., 2016). This plant order has been the focus of several broad phylogenetic studies (Långström & Chase, 2002;Nazaire & Hufford, 2012;Cohen, 2014;Weigend & al., 2014;Luebert & al., 2016). ...
... Boraginales are a group of flowering plants consisting of up to 2700 species of trees, shrubs, or (often) herbs, with a nearly worldwide distribution (Mabberley, 2008;Luebert & al., 2016). This plant order has been the focus of several broad phylogenetic studies (Långström & Chase, 2002;Nazaire & Hufford, 2012;Cohen, 2014;Weigend & al., 2014;Luebert & al., 2016). The Boraginales have been subject to differing circumscriptions over the years, having been treated as equivalent to a single, large family Boraginaceae s.l. ...
... The Boraginales have been subject to differing circumscriptions over the years, having been treated as equivalent to a single, large family Boraginaceae s.l. (e.g., Mabberley, 2008;APG IV, 2016) with several subfamilies, or split into as many as eleven families (e.g., Weigend & al., 2014;Luebert & al., 2016), including the Cordiaceae, Ehretiaceae, Heliotropiaceae, Hydrophyllaceae, and a more narrowly circumscribed Bora gin aceae s.str. We elect to treat the Boraginaceae in this strict sense, as in Luebert & al. (2016) and Chacón & al. (2016), and our use of the name "Boraginaceae" is with this circumscription for the remainder of this article. ...
... The order Boraginales of the Lamiids clade has 125 genera and 2700 species distributed globally (Luebert et al. 2016). The classification of this order has changed over time, especially after the introduction of molecular and phylogenetic studies. ...
... Stevens (2001-onwards) treats the Boraginales as having eight families: Boraginaceae, Codonaceae, Cordiaceae, Ehretiaceae, Heliotropiaceae, Hydrophyllaceae, Namaceae and Wellstediaceae. However, Luebert et al. (2016) based on combined phylogenetic data (Refulio-Rodrıguez & Olmstead 2014;Weigend et al. 2014) and morphological characters favoured recognition of eleven monophyletic families within this order and resolved them into two major clades, namely BOR-I and BOR-II. BOR-I is composed of the families Codonaceae and Wellstediaceae, sister to the cosmopolitan Boraginaceae sensu stricto (s.str.), while BOR-II is composed of families Hydrophyllaceae, Namaceae, Heliotropiaceae, Cordiaceae sensu lato (s.l.) (Cordiaceae, Coldeniaceae, and Hoplestigmataceae) and Ehretiaceae sensu lato (Ehretiaceae and Lennoaceae). ...
... ex Dumort. comprises ∼350 species of trees, shrubs, or rarely lianas (Stevens 2001onwards;Luebert et al. 2016). These species are distributed worldwide, especially in the tropics and subtropics, and consist of two genera (Cordia L. and Varronia P. Browne) according to Luebert et al. (2016) or three genera (Cordia, Varronia and Hoplestigma Pierre) according to Stevens (2001-onwards). ...
The order Boraginales of the Lamiids clade encompasses a diverse group of plants that currently has a global distribution. Its fossil record is scanty. Herein we report a fossil wood of Cordia L. of the family Cordiaceae from the Tipam Sandstone (late Miocene) sediments of Assam, northeast India. This finding not only throws light on the phytogeography and diversification of the Cordiaceae, but also provides valuable insights into the Neogene vegetation of the region.
... The Ehretiaceae (Ehretioideae) is a family of the flowering plant order Boraginales. The family, as most recently circumscribed, contains seven genera (Bourreria, Cortesia, Ehretia, Halgania, Lepidocordia, Rochefortia, and Tiquilia) and comprises more than 150 species widely spread in tropical and subtropical regions [1]. The Ehretiaceae members are mostly trees with the following characteristics: leaves are entire and alternate in arrangement; inflorescence is terminal or axillary; flowers are 5-merous; bisexual or unisexual; corolla is white, blue, or red; shape is tubular, campanulate, or rotate; five stamens, ovary in a slender terminal style, and two stigmas divided slightly or deeply; four ovules in two or four locules; fruit drupaceous, dry or fleshy [1][2][3]. ...
... The family, as most recently circumscribed, contains seven genera (Bourreria, Cortesia, Ehretia, Halgania, Lepidocordia, Rochefortia, and Tiquilia) and comprises more than 150 species widely spread in tropical and subtropical regions [1]. The Ehretiaceae members are mostly trees with the following characteristics: leaves are entire and alternate in arrangement; inflorescence is terminal or axillary; flowers are 5-merous; bisexual or unisexual; corolla is white, blue, or red; shape is tubular, campanulate, or rotate; five stamens, ovary in a slender terminal style, and two stigmas divided slightly or deeply; four ovules in two or four locules; fruit drupaceous, dry or fleshy [1][2][3]. ...
... This classification is also supported by the Angiosperm Phylogeny Group (APG) and several systematic plant studies [10][11][12][13][14][15]. However, different phylogenetic studies in recent decades have recognized Ehretiaceae as a separate family within the order Boraginales [1,[16][17][18][19][20][21]. Most research that has covered the evolutionary relationships of the family Ehretiaceae has used a few genes originating from mitochondrial, chloroplast, and nucleus DNA [22]. ...
Ehretiaceae is a family in the order Boraginales. It contains more than 150 species. The Ehretiaceae classification has remained elusive and changed over time from subfamily to family, or vice versa. In this paper, we sequenced, characterized, and analyzed the complete chloroplast (cp) genomes of Ehretia cymosa and Ehretia obtusifolia, and their cp genomes were compared to those of related species. The length of the chloroplast genomes of E. cymosa was 156,328 bp, whereas that of E. obtusifolia was 155,961 bp. Each genome contained 114 genes, including 80 protein-coding genes, 4 rRNA genes, and 30 tRNA genes. Repeat analysis revealed that complement, forward, palindromic, and reverse repeats were present in the chloroplast genomes of both species. Simple sequence repeat analysis showed that the chloroplast genomes of E. cymosa and E. obtusifolia comprise 141 and 139 microsatellites, respectively. Phylogenetic analysis based on Bayesian and maximum likelihood analyses divided the order Boraginales into two well-supported clades. The first clade includes a single family (Boraginaceae), and the second clade includes three families (Ehretiaceae, Cordiaceae, and Heliotropiaceae). This study provides valuable genomic resources and insights into the evolutionary relationships within Boraginales.
... gov/Genbank), including one published Keraunea sample (Muñoz-Rodríguez et al. 2022). We supplemented these data with eight samples representing families in the Boraginales sensu Luebert et al. (2016). All sequences were retrieved with a custom R script using functions from ape package (Paradis and Schliep 2019). ...
... The phylogenetic realignment of the genus Keraunea as nested within the Ehretiaceae requires a slightly emended description of the family. Keraunea represents the first lianescent member of a family that otherwise includes trees, shrubs and perennial herbs (Luebert et al. 2016). The inflorescence structure of Keraunea is also unique not just within the Ehretiaceae but in the Boraginales as a whole. ...
... The inflorescence structure of Keraunea is also unique not just within the Ehretiaceae but in the Boraginales as a whole. Keraunea does, however, present the only character believed to hold together the Ehretiaceae, bifid styles with two stigmatic branches (Luebert et al. 2016). We provide an emended family description below (see Taxonomic treatment) and a discussion of the unique inflorescence of Keraunea. ...
Keraunea is an enigmatic genus of lianescent shrubs endemic to Brazil and found within the Caatinga and Mata Atlântica phytogeographic regions. When first published, Keraunea was included in the Convolvulaceae and there has been considerable recent disagreement about its true family placement on the Angiosperm tree of life. Based on further assessment of morphology and a new comprehensively-sampled combined phylogenetic analysis of nuclear and plastid genes from recently published DNA sequence data, we settle the position of the genus within the Ehretiaceae as sister to the Australian genus Halgania Gaudich. and provide an expanded family description. We recognize five species within Keraunea,
... Cordiaceae, without exception woody, encounter ca 400 species of worldwide distribution, especially in the tropics and subtropics (Luebert et al., 2016). They belong to the Primarily Woody Boraginales (PWB: Gottschling, 2003) being distinguished by indehiscent fruits and a multilayered endocarp that bursts at predetermined dehiscence lines during germination (Diane et al., 2002;Gottschling, 2004;Heigl et al., 2020). ...
... They belong to the Primarily Woody Boraginales (PWB: Gottschling, 2003) being distinguished by indehiscent fruits and a multilayered endocarp that bursts at predetermined dehiscence lines during germination (Diane et al., 2002;Gottschling, 2004;Heigl et al., 2020). Cordiaceae are a monophyletic group based on molecular phylogenetics and apomorphies such as the four stigmatic branches of the gynoecium, plicate cotyledons and an undivided endocarp (Gottschling et al., 2005;Heigl et al., 2020;Luebert et al., 2016) and comprise Cordia and Varronia P.Browne (Miller & Gottschling, 2007). As inferred from molecular phylogenetics, Cordia segregates into the subclades Sebestena Boehm., Collococcus and Myxa, whereas the latter two are closer related (Gottschling et al., 2005). ...
... Axial placentation is abundant across Boraginales, and Cordiaceae have hemi(ana)tropous (versus anatropous) ovules as probable apomorphy (Heigl et al., 2020). Of the four initial ovules, one through three are frequently abortive and consequently, many fruits of Cordiaceae contain a single seed only (Luebert et al., 2016). ...
Detailed anatomical studies of flowers and fruits may render traits of phylogenetic importance, but are still rare in the Boraginales. An Old World clade of Cordia comprises many dioecious species, but the floral anatomy is largely unexplored and, hence, the structural basis for sex separation. Fresh material of buds, functionally unisexual flowers and fruits were studied using a stack shot camera. Fixed such material of Cordia crenata , C. cf. grandicalyx and C. sinensis was sectioned using a microtome and stained with safranin red and astra blue, and the gynoecial surface was studied using SEM. The stamens of the female flowers are largely intact, but do not produce pollen, whereas the gynoecium of the male flowers is highly reduced though develops ovules lacking an embryo sac. The morphologically rather unreduced stamens of the female flower may have undergone a shift of function from producing pollen towards attracting pollinators by optical signals. The outer parenchyma of the gynoecium is strongly stained and after fertilisation, one through three ovules are maturing towards seeds. The presence of more than a single seed enclosed in the undivided endocarp is rare in Cordiaceae.
... 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). ...
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.
... 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). ...
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.
... Cordiaceae (Cordioideae) is a family within the flowering plant order Boraginales. The Cordiaceae family is split into two genera, Cordia and Varronia, and has over 400 species [1]. The species of the Cordiaceae family are shrubs or trees; the leaves are arranged in a spiral, simple and entire; the flowers are mostly 5-merous, actinomorphic; the petals are white; and the fruit is a drupe, thinly fleshy or dry and hard [2]. ...
... The second clade consists of Cordiaceae, Ehretiaceae, and Heliotropiaceae; Cordiaceae resolved as sister to Ehretiaceae, which is consistent with previous phylogenetic studies [56,87]. Our results support treating the order Boraginales to include several distinct families, consistent with a number of recent molecular studies [1,12,56,88] and contrary to what the APG IV system suggested, which treated the Boraginales to include only one family, Boraginaceae [9]. ...
Cordiaceae is a family comprising more than 400 species in the order Boraginales. The classification of this family has undergone changes over time, transitioning between family and subfamily status. In the present study, the complete chloroplast (cp) genomes of Cordia monoica and Cordia sinensis were sequenced, and their cp genomes were then characterized, analyzed, and compared to those of closely related taxa. The lengths of the cp genomes of C. monoica and C. sinensis were 151,813 bp and 152,050 bp, respectively. Both genomes consisted of 114 genes, divided into 4 ribosomal RNA genes, 30 transfer RNA genes, and 80 protein-coding genes. We observed a unique gene inversion in the trnM-rbcL region of both Cordia species. The long repeats analysis revealed that both species’ chloroplast genomes contained forward and palindromic repeats. The simple sequence repeats (SSRs) analysis detected 155 microsatellites in each genome, with the majority being mononucleotide repeats (A/T). Phylogenetic analysis based on maximum likelihood and Bayesian analyses confirmed two major clades in the order Boraginales: clade I comprised Boraginaceae, while clade II included Cordiaceae, Ehretiaceae, and Heliotropiaceae. This study expands our knowledge of the evolutionary relationships across the order Boraginales and offers useful genetic resources.
... Ehretiaceae has been delimited in different ways since it was resurrected from synonymy within a widely defined Boraginaceae. In this paper we follow the delimitation of the familial classification of the Boraginales by the 20 members of the Boraginales Working Group which recognises 11 families (Luebert et al. 2016) rather than that of Gottschling et al. (2016) in which Cordiaceae and Hoplestigmataceae are included in Ehretiaceae. ...
... Placement within Ehretiaceae (Boraginales) has is also confirmed by morphological evidence. Initially, this seemed surprising, since the climbing habit and presence of a bract is not included in the morphological circumscription of the family (Luebert et al. 2016). However, in the more extensive treatment of Ehretiaceae by Gottschling et al. (2016) (albeit with the inclusion of Cordiaceae and Hoplestigmataceae), it is reported that some species of Rochefortia Sw. and Bourreria P. Browne are also scandent or lianescent, and a bract is reported in Cortesia (Mirra et al. 2022), here resolved as one of the possible sister genera to Keraunea (Fig. 2). ...
We definitively place Keraunea , a genus of showy forest climbers from remnants of the Mata Atlântica of Brazil, in Ehretiaceae. Previously Keraunea had been ascribed to Convolvulaceae based on morphology, or divided between Malpighiaceae and Ehretiaceae based on molecular analyses (polyphyletic). Keraunea is morphologically anomalous in the Ehretiaceae, having fruits which are held in the centre of a large wing-like bract by adnation of the pedicel, and due the stem-twining habit of some species. However, morphologically Keraunea shares two apomorphies with monotypic Cortesia Cav., halophytic shrubs of Argentina: 1) “two parted endocarps” (the fruit contains two endocarps each with two 1-seeded cells, while four 1-seeded endocarps are plesiomorphic in Ehretiaceae) and 2) a large bract that subtends the flower (absence of bracts is plesiomorphic in Ehretiaceae). A combined tree using four markers (ITS, trn L-F, rbc L and mat K) places the three species of Keraunea sampled unambiguously in a clade with Ehretia + Cortesia + Halgania and monophyly for Keraunea is shown with high support. In an ITS tree Keraunea is sister to Cortesia with low support.
We expand the generic description of Keraunea including the first account of the internal fruit structures and the seed, and present new data on the vegetative architecture including variation within the genus, some species being stem twiners while one species at least, is described as a scandent shrub. Keraunea lombardiana , previously included within K. brasiliensis , is formally described from Minas Gerais and Bahia as the third species of the genus and both these species are described. In all five species are recognised but two, known to us only from digital images, are not formally described because sufficiently detailed descriptions cannot be made. All five species are mapped, and provisional conservation assessments are recorded, of either Endangered or Critically Endangered. The state of Bahia, with three species, has the highest species diversity, mainly in dry forest. Three species appear confined to moist coastal forest, extending southwards from Bahia to the state of Rio de Janeiro.
Keraunea remains incompletely known. Not one of the species has both anthetic flowers and fruits described. Studies on pollen, germination, anatomy, embryology and phytochemisty are entirely lacking. Field observations of pollination, seed dispersal and phenology are also required. However, the most urgent requirement is undoubtedly a formal taxonomic revision based on a full herbarium search and targeted fieldwork, with full IUCN 2012 extinction risk assessments for each taxon. This is urgent because it seems that all the taxa that we present in this paper (and more that can be expected to be found) appear to be highly range restricted and generally not to occur in protected areas, and so appear to be highly threatened.
... However, the taxonomic treatment of Boraginales was revised with the development of molecular systematics. Eleven families were adopted in Boraginales based on molecular evidence [2][3][4]. The four subfamilies (Boraginoideae, Ehretioideae, Cordioideae, and Heliotropioideae) form independent families (Boraginaceae, Ehretiaceae, Cordiaceae, and Heliotropiaceae). ...
... The four families had identical tree topologies in the phylogenetic results. It is difficult to say whether Boraginales should be treated as one family, as APG does, or if it should be split into many sections [2,8]. According to the definition of monophyly, these two views are strongly supported in this study. ...
The Boraginales (Boraginaceae a.l.) comprise more than 2450 species worldwide. However, little knowledge exists of the characteristics of the complete plastid genome. In this study, three new sequences representing the first pt genome of Heliotropiaceae and Cordiaceae were assembled and compared with other Boraginales species. The pt genome sizes of Cordia dichotoma, Heliotropium arborescens, and Tournefortia montana were 151,990 bp, 156,243 bp, and 155,891 bp, respectively. Multiple optimal codons were identified, which may provide meaningful information for enhancing the gene expression of Boraginales species. Furthermore, codon usage bias analyses revealed that natural selection and other factors may dominate codon usage patterns in the Boraginales species. The boundaries of the IR/LSC and IR/SSC regions were significantly different, and we also found a signal of obvious IR region expansion in the pt genome of Nonea vesicaria and Arnebia euchroma. Genes with high nucleic acid diversity (pi) values were also calculated, which may be used as potential DNA barcodes to investigate the phylogenetic relationships in Boraginales. psaI, rpl33, rpl36, and rps19 were found to be under positive selection, and these genes play an important role in our understanding of the adaptive evolution of the Boraginales species. Phylogenetic analyses implied that Boraginales can be divided into two groups. The existence of two subfamilies (Lithospermeae and Boragineae) in Boraginaceae is also strongly supported. Our study provides valuable information on pt genome evolution and phylogenetic relationships in the Boraginales species.
... 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]. ...
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.
... The RSA-POM herbarium was used as a reference to help verify plant identifications, along with the Jepson eFlora (Jepson Flora Project 2019) and the Intermountain Flora (Cronquist et al. 1972(Cronquist et al. -2017. The classification follows the Jepson Flora Project (2019), except for Boraginales, which follows Luebert et al. (2016) and Cryptantha, which follows Hasenstab-Lehman et al. Vegetation.-A list of major vegetation types was compiled during the first collecting season. ...
... If a historical specimen is cited and that taxon was not relocated during the study, the herbarium where the specimen is housed is indicated if not RSA-POM. The classification follows the order classification of Angiosperm Phylogeny Group IV (Stevens 2001 onwards), familial classification of Jepson eFlora (Jepson Flora Project 2019), other than Boraginaceae, for which an updated treatment was utilized (Luebert et al. 2016) and Cryptantha which follows Hasenstab-Lehman (2012). Non-native taxa are designated by an asterisk (*), CNPS-listed rare taxa are shown with a dagger ( †) and taxa previously found but not relocated during this study are indicated with a diamond (♦). ...
Adobe Valley and the Adobe Hills lie east of the Sierra Nevada in Mono County, California, and are within the Great Basin Floristic Province. The flora of Adobe Valley and the surrounding hills is influenced by the Great Basin Desert to the east, the Sierra Nevada to the west and the northern Mojave Desert to the south. Adobe Valley is surrounded by the Adobe Hills, Benton Range and Glass Mountain region, creating a circular closed basin with many small tributaries feeding into it. This topography contributes to the creation of a rare wetland complex, including alkali meadows, marshes, and lakes at the base of the hills, which represent some of the more botanically interesting terrain in the region. Prior to this study, the wetland complex of Adobe Valley had not been systematically investigated with regard to its botanical diversity. The vegetation types associated with the wetland complex are listed as threatened by the California Natural Diversity Database and face a number of conservation concerns including water pumping, overgrazing, and possible wind energy development. A total of 1525 herbarium specimens were collected within the study area over 59 days in the field from 2016 to 2018. The vascular flora of the Adobe Valley and Hills includes 397 minimum-rank vascular plant taxa, representing 194 genera and 61 families. Six taxa are only known from historical collections, 21 taxa are non-native and 27 taxa have conservation status. The results of the inventory are presented here in an annotated checklist, along with descriptions of vegetation alliances.
... 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. ...
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.
... The systematic order and taxonomic circumscription of the families follow the classification proposed by PPG I (2016) for lycophytes, ferns and fern allies, by Christenhusz et al. (2011) for gymnosperms and by APG IV (2016) for angiosperms, with the exception of Dipsacales (Reveal 2011), Caryophyllales (Hernández-Ledesma et al. 2015, Boraginales (Luebert et al. 2016) and Celastrales (Simmons et al. 2023). The disputed family Chenopodiaceae was included in Amaranthaceae based on the recent molecular study by Morales-Briones et al. (2020). ...
Critical species inventories provide primary biodiversity data crucial for biogeographical, ecological, and conservation studies. After six years, a second update to the inventory of the vascular flora native to Italy is presented. It provides details on the occurrence at regional level and, for the first time, floristic data for San Marino. The checklist includes 8,241 species and subspecies, distributed in 1,111 genera and 153 families; 23 taxa are lycophytes, 108 ferns and fern allies, 30 gymnosperms, and 8,080 angiosperms. The species/subspecies endemic to Italy are 1,702, grouped in 71 families and 312 genera. The taxa currently occurring in Italy are 7,591, while 545 taxa have not been confirmed in recent times, 94 are doubtfully occurring in the country, 11 are data deficient, and 236 are reported by mistake and to be excluded at national level. Out of the 545 not confirmed taxa, 28 are considered extinct or possibly extinct.
... Most species, typically grow in alluvial deposits and along margins of fluvial streams (Taylor 2012). The genus has been placed in the Boraginaceae and Hydrophyllaceae at different times, but more recently, it is recognized in the Namaceae according APG IV (Luebert et al. 2016, Molinari-Novoa 2016. Hitchcock´s (1933a, b) Nama monograph included only 32 taxa, but approximately some 28 other taxa were described in the following 80 years (Taylor 2012). ...
Background:
Although most of the Nama taxa grow on sandy substrates, none of them is restricted to the coastal dune environment. A small population of a unique looking taxon was found on the Pacific coast of the Baja California peninsula, which is here presented as a new species.
Question and/or Hypothesis:
In the morphological analysis in the individuals of a putative new taxon, some peculiarities stand out, such as a suffruticose herbaceous perennial growth form with erect and compact branching, featuring a small cushion, unlike the rest of the known species, which are rather procumbent, or erect in form. It also has revolute leaf margins in the blade.
Studied Species:
Nama asuncionensis sp. nov. (Namaceae) and morphologically similar species.
Study Site:
Central Pacific coast of Baja California, Mexico.
Method:
Through bibliographic consultation, review of herbarium specimens, and photographs, a comparative morphological analysis was carried out with other Nama species of the peninsula and mainland Mexico.
Results:
Based on the comparative analyses, Nama asuncionensis is here presented and illustrated as a new species.
Conclusions:
Nama asuncionensis is a micro-endemic species restricted to the Pacific coastal environment of the peninsula. We propose that this new taxon be included in the Eunama section of the genus based on its morphological characteristics, including a membranous loculicidal dehiscent capsule, free styles, and sepals divided to the base. This taxon seems to be limited to a narrow coastal strip, with high relative humidity, where it develops its fresh leaves and flowers during the mild season.
... The Boraginaceae family, which ranks ninth among the families with the highest number of species in Turkey, includes 357 taxa, 34 genera, 325 species, 16 subspecies, and 16 varieties. Onosma L., the largest genus of the Boraginaceae family, is represented by a total of 150 species worldwide [31,32]. In Turkey, Onosma L. is represented by 97 species, four varieties, and one hybrid species, of which 50 species and one type are endemic [33,34]. ...
The aboveground and root parts of Onosma mutabilis were extracted using subcritical water and the process was optimized with response surface methodology. The composition of the extracts was determined by chromatographic methods and compared to that of conventional maceration of the plant. The optimum total phenolic contents for the aboveground part and the roots were 193.9 and 174.4 µg/g, respectively. These results were achieved at a subcritical water temperature of 150 • C, an extraction time of 180 min, and a water/plant ratio of 0.1, for both parts of the plant. Principal component analysis revealed that the roots contained mainly phenols, ketones, and diols, with the aboveground part mostly alkenes and pyrazines, whereas the extract from maceration contained mainly terpenes, esters, furans, and organic acids. The quantification of selected phenolic substances showed that subcritical water extraction compared favorably to maceration, especially with respect to pyrocatechol (1062 as compared to 10.2 µg/g) and epicatechin (1109 as compared to 23.4 µg/g). Furthermore, the roots of the plant contained twice as much of these two phenolics compared to the aboveground part. Subcritical water extraction of O. mutabilis is an environmentally friendly method that can extract selected phenolics at higher concentrations compared to maceration.
... Onosma species belongs to the Boraginaceae family. The Boraginaceae family contains more than 100 genera and over 200 species, which are classified into five subfamilies: Boraginoideae, Cordioideae, Ehretioideae, Hydrophylloideae, and Lennooideae [30,31] (Table 1). ...
The genus Onosma belongs to the Boraginaceae family and contains over 230 species. The present review sheds light on the ethnopharmacology, phytoconstituents, bioactivity, and toxicology of the Onosma species from previous investigations. Furthermore, the paper also highlights the unresolved issues for the future investigations. The review included previous studies of the genus Onosma available from Google Scholar and Baidu Scholar, Science Direct, SciFinder, Wiley Online Library, and Web of Science. Until now, more than 200 chemical compounds have been detected from the genus Onosma, including naphthoquinone (33), flavonoids (30), hydrocarbon (23), phenolic (22), ester (17), alkaloids (20), aromatics (12), carboxylic acid (11), fatty acids (9), terpenoids (10), while the most important ones are rosmarinic, ferulic, protocatechuic, chlorogenic, caffeic, p-coumaric acids, and apigenin. The Onosma species are reported as traditional medicine for wound healing, heart disease, and kidney disorders, while the pharmacological investigations revealed that the extracts and the phytochemicals of Onosma species have different therapeutic properties including antioxidant, enzyme inhibitory, antitumor, hepatoprotective, antiviral, anti-inflammatory, and antimicrobial actions. The summarized knowledge in this review provides valuable ideas for the current and future drug discovery and a motivation for further investigation on the genus Onosma.
... 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). ...
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
... 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. ...
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.
... 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). ...
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.
... 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. ...
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. ...
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.
... 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. ...
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.
... 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. ...
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.
Critical and updated alien species inventories are crucial to properly manage and contrast biological invasions. After six years, a second update to the inventory of the vascular flora alien to Italy is presented. It provides details on the occurrence at administrative regional level and, for the first time, floristic data for San Marino. The checklist includes 1,782 taxa (1,673 species, 28 subspecies, 69 hybrids, and 12 cultivar groups not attributable to a specific binomial), distributed in 786 genera and 159 families; 2 taxa are lycophytes, 13 ferns and fern allies, 37 gymnosperms, and 1,730 angiosperms. Among these, 154 taxa are archaeophytes and 1,628 neophytes. The alien taxa currently established in Italy are 899 (649 naturalized and 250 invasive), while 796 taxa are casual aliens, 1 are not assessed, 38 have not been confirmed in recent times, 4 are considered extinct or possibly extinct in the country, 40 are doubtfully occurring in Italy, 4 are of unknown regional distribution, and 110 are reported by mistake and to be excluded at national level. This checklist allows to establish an up-to-date number (10,023) of taxa currently constituting the whole (either native or alien) Italian flora.
Euploca wightiana P.Javad & Ancy sp. nov., a new species of the family Heliotropiaceae is described from the Coimbatore district of South India. Morphologically, it is close to Euploca marifolia but differs in having erect (versus prostrate) habit, stem woody throughout (versus stem woody only at base) with dense hairs (versus sparsely hairy), oblong leaves (versus lanceolate) with obtuse apex (versus acute), inflorescence with distantly placed flowers (versus inflorescence with closely placed flowers), a 0.6 mm long cylindrical style (versus style less than 0.3 mm long), a unifid, capitate stigma (versus bifid, conical) and densely hirsute nutlets (versus minutely hairy or glabrous). Detailed descriptions, photo plates, and distribution maps are provided. Differences from morphologically allied congeners are discussed. The status of the new species is provisionally assessed here as ‘critically endangered (CR)' following the IUCN red list categories and criteria.
Angiosperms (flowering plants) are by far the most diverse land plant group with over 300,000 species. The sudden appearance of diverse angiosperms in the fossil record was referred to by Darwin as the “abominable mystery,” hence contributing to the heightened interest in angiosperm evolution. Angiosperms display wide ranges of morphological, physiological, and ecological characters, some of which have probably influenced their species richness. The evolutionary analyses of these characteristics help to address questions of angiosperm diversification and require well resolved phylogeny. Following the great successes of phylogenetic analyses using plastid sequences, dozens to thousands of nuclear genes from next‐generation sequencing have been used in angiosperm phylogenomic analyses, providing well resolved phylogenies and new insights into the evolution of angiosperms. In this review we focus on recent nuclear phylogenomic analyses of large angiosperm clades, orders, families, and subdivisions of some families and provide a summarized Nuclear Phylogenetic Tree of Angiosperm Families. The newly established nuclear phylogenetic relationships are highlighted and compared with previous phylogenetic results. The sequenced genomes of Amborella, Nymphaea, Chloranthus, Ceratophyllum, and species of monocots, Magnoliids, and basal eudicots, have facilitated the phylogenomics of relationships among five major angiosperms clades. All but one of the 64 angiosperm orders were included in nuclear phylogenomics with well resolved relationships except the placements of several orders. Most families have been included with robust and highly supported placements, especially for relationships within several large and important orders and families. Additionally, we examine the divergence time estimation and biogeographic analyses of angiosperm on the basis of the nuclear phylogenomic frameworks and discuss the differences compared with previous analyses. Furthermore, we discuss the implications of nuclear phylogenomic analyses on ancestral reconstruction of morphological, physiological, and ecological characters of angiosperm groups, limitations of current nuclear phylogenomic studies, and the taxa that require future attention.
Eriodictyon capitatum (Namaceae) is a narrowly distributed shrub endemic to western Santa Barbara County, where it is known from only 10 extant California Natural Diversity Database element occurrences (EOs). Owing to low numbers of plants in nature, a limited overall extent, and multiple current threats, E. capitatum is listed as Endangered under the Federal Endangered Species Act and as Rare under the California Native Plant Protection Act. In the present study, high-throughput DNA sequence data were analyzed to investigate genetic diversity within and among all accessible EOs; to determine the extent of genetic isolation among EOs; to examine clonality within EOs; and to examine the taxonomic circumscriptions of E. capitatum, E. altissimum, E. angustifolium, and E. californicum through phylogenomic analysis. Population genetic analyses of E. capitatum reveal a pattern of strong genetic differentiation by location/EO. The clonality assessment shows that certain small EOs may support relatively few multilocus genotypes. The phylogenomic analyses strongly support the present-day taxonomic circumscriptions of both E. altissimum and E. capitatum, showing them to be reciprocally monophyletic and sister with strong support. Taken together, these results paint a picture of an evolutionarily and morphologically distinct species known from relatively few, genetically isolated stations.
Background
The systematics of family Boraginaceae draw attention of many botanists for many years. The current study's primary goals are to clarify phenetic and phylogenetic relationships within Boraginaceae according to morphology and molecular characteristics and to evaluate the morphological characters that can be applied in systematics of Boraginaceae.
Results
The macromorphological characters of 39 species, 2 subspecies and 5 varieties of wild boraginaceous plants were extracted and subjected to phenetic and principal component analysis that was performed for detecting the most important characters differentiating the studied taxa. The generated dendrogram is divided into five clear groups; Arnebia decumbens var. macrocalyx and Heliotropium curassavicum are the most distantly related species, while Echium angustifolium subsp. angustifolium and E. angustifolium subsp. sericeum are the most closely related species. The phylogenetic relations among the examined taxa were determined using DNA barcoding of the rbcl gene. The phylogenetic analysis generated a cladogram showing that among the studied taxa of Boraginaceae there is a bolster for three clear lineages with resolved relationships.
Conclusions
It is concluded that the chosen morphological characters were important in species delimitation, where more than half of the total morphological variations (67.94%) were explained by the first two principal components, indicating that the morphological characters showed high variability, which is useful for discrimination, and these characters, in addition to molecular characters, shared in drawing the phenetic and phylogenetic relationships within Boraginaceae that were considered not monophyletic groups. Boraginaceae contained some monophyletic genera such as Heliotropium and Alkanna , while the other studied taxa expressed a non-monophyletic relationships.
Recent phylogenetic studies have challenged the traditional classification of the subtribe Cynoglossinae, which was based on nutlet characteristics. To investigate the evolution of fruit traits related to dispersal modes in this complex group, we conducted a study using scanning electron microscopy to examine 28 taxa representing all previously recognized lineages of the subtribe. Cynoglossinae displays four main types of nutlets: marginate, emarginate, flat wing, and incurved wing. Our findings reveal the arrangement of glochids and their structure, including the number of apical hooks, and their surface ornamentation is highly variable both across and within these four main types. We reconstructed the phylogenetic tree with four main clades (i.e., Clades I–IV) using Maximum likelihood analysis. In addition, we mapped four nutlet traits (i.e., nutlet margin, nutlet glochid, glochid sculpture, and glochid hooks) associated with different dispersal modes using ancestral state reconstruction analysis. The ancestor of subtribe Cynoglossinae was inferred to have marginate and glochidiate nutlets, with glochids externally provided with complex spiny tubercles and bearing at least four hooks at the tip as well as epizoochory dispersal mode. These ancestral states have been retained in Clade II (East Asian-Australian-African Cynoglossum-Lindelofia clade) and Clade III (Mediterranean-Asian Cynoglossum-Solenanthus-Trachelanthus clade), with several independent shifts to emarginate fruits. Transitions from marginate to wide-winged nutlets have occurred at least four independent times, facilitating wind-assisted dispersal in Clade I (Microparacaryum clade), Clade IV (Paracaryum-Mattiastrum-Rindera clade), Rindera tetraspis, and Mattiastrum crista-galli (both belonging to Clade III), where a transition from tuberculate to papillate surface has also occurred.
A synopsis of Heliotropiaceae in Thailand is presented and, as part of this, the taxonomic history, identification keys, distribution maps, herbarium specimen citations and diagnostic characters are provided. Two genera and 12 species are recognised and four new combinations are proposed of Tournefortia under the genus Heliotropium , namely H. hookeri , H. intonsum , H. montanum and H. ovatum . Two species are newly recorded in Thailand: Euploca ovalifolia and H. hookeri . The neotype and lectotype are designated here for Tournefortia montana and T. boniana , respectively.
Ehretia pranomiana sp. nov. is illustrated and described from northern and south-western Thailand. The new species is similar to E. macrophylla and E. acuminata by having elliptic leaves with a serrate margin, but can be distinguished by being a scrambling shrub with densely hispid leaf blades, corymbose inflorescence with 4–10 flowers, corolla with erect lobes and calyx longer than fruit at maturity. An updated identification key to the species of Ehretia known from Thailand is provided.
A synopsis of Heliotropiaceae in Thailand is presented together with the taxonomic history, identification keys, distribution, phenology, herbarium specimen citations and diagnostic characters of each species. The taxonomic character of the family is of terminal style with a conical stigma structure forming a basal ring around the style. There are 12 species: five species each in Euploca and Tournefortia and two species in Heliotropium . Three species are newly recorded in Thailand, Euploca ovalifolia , Tournefortia hookeri and T. montana . Lectotypes are selected for T. boniana and T. tetrandra var. angustifolia.
The genus Keraunea was recently described in the Convolvulaceae Juss. family. Two species are currently recognised, both from Brazil. Molecular sequence data using three commonly applied DNA markers (matK, rbcL and the nuclear ribosomal Internal
Transcribed Spacer) show that neither species is correctly placed in Convolvulaceae but indicates that the type, K. brasiliensis, should be placed in Malpighiaceae despite several morphological anomalies. The second species, K. capixaba, should be placed in Ehretiaceae. Given the surprising nature of these results, further studies are recommended before formal reclassification of these two taxa is made.
Pontechium maculatum (Boraginaceae) is a species of high conservation concern in the Bulgarian flora. It is assigned a national IUCN category ‘Vulnerable’, legally protected according to the national Biological Diversity Act, and listed in Annex II of the Habitats Directive (Council Directive 92/43/EEC). Recent studies in one of the largest populations in the country revealed some new data about the species which are reported here. The population studied is situated in Mt Lozenska, Mt Sredna Gora (Western) floristic region. Relevant taxonomic literature states that P. maculatum is a biennial plant, however, our observations revealed the plants in Mt Lozenska are perennial (possibly short-lived). A great variation in the corolla-colour was observed, most plants having bluish to violet corolla and only a few – reddish to dark red. This can be explained by either natural variation in the population, not reported in the taxonomic literature so far, or by hybridisation with Echium vulgare which cooccurs in the same locality. Large fluctuation in the number of flowering plants in two consecutive years was recorded, with some 180 blooming specimens in 2019 and only about 25 in 2020. Studies in the genome size of this population revealed the plants have significantly higher DNA-content (1C-value) than plants from the typical populations of the species in Northeast Bulgaria floristic region, e.g. Kabiyushka Mogila locality. Further biosystematics studies(chromosome number, genome size, genetic studies, etc.) are currently in progress to explain the observed phenomena within the Bulgarian populations of P. maculatum.
Lompoc yerba santa Eriodictyon capitatum (Namaceae) is an evergreen shrub endemic to west Santa Barbara County in coastal southern Calif. The species was listed as rare under the Calif. Endangered Species Act in 1979 and as endangered under the U.S. Endangered Species Act in 2000. At Federal listing in 2000, Lompoc yerba santa was known from four locations (five occurrences) in three geographic areas. In 2022, the species is known from 11 occurrences in four geographic populations of which 10 occurrences are considered extant and 1 occurrence likely extirpated: Orcutt Oil Field in the Solomon Hills, 2 occurrences; Burton Mesa on Vandenberg Space Force Base North, 6 occurrences; La Salle Canyon on Vandenberg Space Force Base South, 1 occurrence; and Santa Ynez Mountains, 2 occurrences. Lompoc yerba santa occurs mostly in chaparral and bishop pine (Pinus muricata) forest, and most occurrences are in sandy soil. The species stands up to 5 m tall and with vertical stems up to 10 cm diameter. Much of its reproduction appears to be vegetative by rhizome production, suggesting low dispersal ability, and it is a habitat specialist with apparent dependence on substrate disturbance and fire. Using all available information and international standards with a precautionary but realistic attitude to uncertainty, Lompoc yerba santa in 2022 meets IUCN criteria for endangered status: the small extent of occurrence is 433 km2 (<5,000 km2) and the small area of occupancy is 2.30 km2 (<10 km2), along with the geographic range being severely fragmented, and the quality of habitat is observed/inferred/projected to be in continuing decline (altered fire regime/fire management practices, climate change with severe drought and increased temperatures).
Nine new combinations including two new names in South American species of Heliotropium (Heliotropiaceae) are proposed in this work: H. antioquianum J.I.M.Melo, replacement name; H. brevilobatum (K.Krause) J.I.M.Melo, comb. nov.; H. fuliginosum (Kunth) J.I.M.Melo, comb. nov.; H. gigantifolium (Killip ex J.S.Mill.) J.I.M.Melo, comb. nov.; H. macrostachyum (Rusby) J.I.M.Melo, comb. nov.; H. rollotii (Killip) J.I.M.Melo, comb. nov.; H. romeroi (I.M.Johnst.) J.I.M.Melo, comb. nov.; H. scabridum (Kunth) J.I.M.Melo, comb. nov., and H. stapfianum J.I.M.Melo, replacement name. Furthermore, Heliotropium romeroi is recorded for the first time in Ecuador and lectotypes are provided for Tournefortia brevilobata K.Krause, T. fuliginosa Kunth, T. macrostachya Rusby, and T. scabrida Kunth.
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.
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.
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.
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.
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.
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.
Aim of study: Aim was to determine the vascular flora of the Ormanüstü Planning Unit (Karadeniz Technical University Education-Research and Application Forest).
Area of study: The research area is under the influence of the Euro-Siberian phytogeographical region and is represented with a high rate (44%, 288 taxa). Irano-Turanian and Mediterranean elements are represented by 26 (4%) and 22 taxa (3%), respectively. Totally 320 taxa (49%) are multi-regional or of unknown origin. In addition, 33 endemic (5%), 12 rare (1.8%), 5 alien (0.8%) and 17 cosmopolitan (2.6%) taxa were identified.
Material and methods: The entire study area was visited periodically and plant samples were collected from 166 sample plots (1000 m2), and then they were pressed with wooden frames. Approximately 1800 vascular plant specimens were collected between the years 2005 and 2008 to form the floristic list.
Main results: Studies revealed the presence of 656 taxa (585 species, 62 subspecies and 9 varieties) belonging to 95 families and 354 genera. Of these, 22 taxa (3.4%) are ferns and 634 (96.6%) are seed plants. The best-represented families are as follows; ;Asteraceae (75), Poaceae (48) and Fabaceae (41). According to richest genera; Campanula (14), Carex (10), Alchemilla (9) and Geranium (9) have the greatest number of taxa.
Highlights: It was concluded that the study area is important in terms of plant diversity. The discovery of a new species from this area, described by us and published in 2009 (Astragalus ansinii Uzun, Terzioğlu & Pal.-Uzun), supports this fact
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
A revised and updated classification for the families of flowering plants is provided. Many recent studies have yielded increasingly detailed evidence for the positions of formerly unplaced families, resulting in a number of newly adopted orders, including Amborellales, Berberidopsidales, Bruniales, Buxales, Chloranthales, Escalloniales, Huerteales, Nymphaeales, Paracryphiales, Petrosaviales, Picramniales, Trochodendrales, Vitales and Zygophyllales. A number of previously unplaced genera and families are included here in orders, greatly reducing the number of unplaced taxa; these include Hydatellaceae (Nymphaeales), Haptanthaceae (Buxales), Peridiscaceae (Saxifragales), Huaceae (Oxalidales), Centroplacaceae and Rafflesiaceae (both Malpighiales), Aphloiaceae, Geissolomataceae and Strasburgeriaceae (all Crossosomatales), Picramniaceae (Picramniales), Dipentodontaceae and Gerrardinaceae (both Huerteales), Cytinaceae (Malvales), Balanophoraceae (Santalales), Mitrastemonaceae (Ericales) and Boraginaceae (now at least known to be a member of lamiid clade). Newly segregated families for genera previously understood to be in other APG-recognized families include Petermanniaceae (Liliales), Calophyllaceae (Malpighiales), Capparaceae and Cleomaceae (both Brassicales), Schoepfiaceae (Santalales), Anacampserotaceae, Limeaceae, Lophiocarpaceae, Montiaceae and Talinaceae (all Caryophyllales) and Linderniaceae and Thomandersiaceae (both Lamiales). Use of bracketed families is abandoned because of its unpopularity, and in most cases the broader circumscriptions are retained; these include Amaryllidaceae, Asparagaceace and Xanthorrheaceae (all Asparagales), Passifloraceae (Malpighiales), Primulaceae (Ericales) and several other smaller families. Separate papers in this same volume deal with a new linear order for APG, subfamilial names that can be used for more accurate communication in Amaryllidaceae s.l., Asparagaceace s.l. and Xanthorrheaceae s.l. (all Asparagales) and a formal supraordinal classification for the flowering plants.
RESUMEN México ha sido considerado un centro de diversidad del género Cordia, del cual se restauró el género Varronia P. Browne (Boraginaceae) que involucra a un grupo importante de especies arbustivas de nuestra fl ora. Esto nos motivo a actualizar la nomenclatura y difundir los cambios propuestos por Miller (2007). Esta revi-sión está basada en la colección del herbario MEXU, pues cuenta con una amplia representación a nivel nacional. Se establece la distribución de 16 especies del género Varronia, la mayoría endémicas y de distribución muy restringida. Para cada una de las especies se establecen los tipos nomenclaturales y sinonimias. Se hacen comentarios acerca de la distribución y hábitat, se incluye una clave para facilitar su identifi cación.
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.
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.
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.
In contrast to previous descriptions, which only state the bicarpellate, syncarpous gynoecium of Wigandia caracasana is superior, this paper demonstrates that it is half inferior and zygomorphic. The abaxial carpel is initiated before the adaxial and constitutes the major part of the ovary. The ovarial chamber is divided at the bottom by both a cross zone and false septa, and at the top by apical septa. The dehiscence tissue of the loculicidally opening capsule develops through the median carpel bundles and the small false septa. The sclerified endocarp, consisting of both inner epidermis and adjacent subepidermal layer, acts as opening tissue. From an entire circular stele that enters the receptacle, ten vascular strands originate from ten gaps. They alternatingly diverge either into the dorsal bundle of a sepal and the single trace of the adjacent stamen, or into the single petal trace and two branches supplying adjacent sepals as marginal bundles. The stele itself vascularizes the gynoecium. Half inferior ovaries seem to be more frequent in the Hydrophyllaceae than previously assumed.
This volume covers the orders Boraginales, Garryales and Solanales (except Convolvulaceae) of the Lamiids (Asterids I) as well as three unplaced families of that clade, i.e. Vahliaceae, Icacinaceae and Metteniusaceae, and the orders Aquifoliales, Escalloniales, Bruniales, Dipsacales and Paracryphiales of the Campanulids (Asterids II). It is the first of two final volumes to (almost) complete the treatment of the Asterids, which started with Vol. VI (Cornales, Ericales, 2004) and continued with Vol. VII (Lamiales, 2004) and Vol. VIII (Asterales, 2007). The present volume provides descriptions for 35 families and altogether 340 genera, including three genera of somewhat uncertain family affiliation. It provides identification keys for families within orders and for all genera within families, and also discusses probable phylogenetic relationships. The wealth of information contained in this volume makes it an indispensable source for all those working in pure and applied plant sciences.
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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.
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.
Five species are here recognized in Wellstedia: W. socotrana on Socotra, W. somalensis sp. nov. and W. laciniata sp. nov. in Somalia, W. filtuensis in Ethiopia, and W. dinteri in Namibia and the adjacent Cape Province. A preliminary phylogenetic analysis suggests that W. dinteri is the sister group to the rest of the genus, while W. filtuensis, W. somalensis and W. socotrana are vicarious sister species. Wellstedia has usually been treated either as a separate subfamily within the Boraginaceae or as a family of its own close to Boraginaceae. Its most notable feature is the capsular fruits (here shown to be hygrochastic), capsules otherwise being unknown in Boraginaceae. It is suggested that the nearest relative of Wellstedia may be Tiquilia in Boraginaceae subfamily Ehretioideae.
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.
The new species Wellstedia robusta, from mountain ridges and slopes south-west of Qandala in north-eastern Somalia, is described and illustrated.
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.
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.
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.
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.
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.
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.
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.
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.
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, ●●, ●●–●●.
The phylogenetic relationships of Mertensia (Boraginaceae), which comprises approximately 45 species in both Asia and North America, have been uncertain, and taxonomists have placed the genus in various tribes of subfamily Boraginoideae, with the most recent placements in Trigonotideae and Cynoglosseae. Our study applies molecular phylogenetic methods to test the monophyly and relationships of Mertensia. We used DNA sequence data from the nuclear ribosomal nrlTS region and four cpDNA regions (matK, ndhF, rbcL, trnL-trnF) to examine the placement of Mertensia among a sampling of accessions from approximately 70% of the genera of Boraginaceae s. 1. Phylogeny reconstructions using maximum parsimony, maximum likelihood, and Bayesian inference were largely congruent with previous molecular phylogenetic analyses of Boraginaceae that had applied far fewer taxa. We recovered five deep clades that correspond to Boraginaceae subfamilies Boraginoideae, Cordioideae, Heliotropioideae, Hydrophylloideae, and Ehretioideae (including Lennoa and Pholisma). In subfamily Boraginoideae, we recovered clades that correspond to the tribes Echiochilieae, Lithospermeae, Cynoglosseae, and Boragineae, although several tribes previously circumscribed on the basis of morphological data were not recovered as monophyletic in our results. Based on the sister relationship between the genus Codon and subfamily Boraginoideae found in our phylogeny reconstructions, we propose Codoneae as a new tribe of Boraginoideae. We recovered strong support for the monophyly of Mertensia and the placement of the monotypic genus Asperugo as its sister. Mertensia and Asperugo were strongly supported as members of Cynoglosseae.