Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications

Ovule and seed structure in Euphorbioideae, one of the five euphorbiaceous subfamilies, is surveyed to evaluate its systematic implications on the basis of 79 species representing four of five tribes. All Euphorbioideae, like two other "uniovulate" subfamilies Acalyphoideae and Crotonoideae, but unlike most of two "biovulate" subfamilies Oldfieldioideae and Phyllanthoideae, consistently have a persistent and palisadal exotegmen composed of radially elongate, sclerotic, and pitted cells. Within Euphorbioideae, the tribe Stomatocalyceae (also with the palisadal exotegmen) is unusual in having vascular bundles in outer integument and clearly distinct from the remaining Euphorbioideae and the other "uniovulate" subfamilies. With the exclusion of Stomatocalyceae, Euphorbioideae are not anatomically divided into major groups such as a pseudanthial and a non-pseudanthial clade, but instead have some remarkable diversity within a tribe, a subtribe, and even a genus in the three ovule and seed characters: (1) the thickness of the inner integument, (2) the thickness of the outer integument, and (3) the presence or absence of an aril. Groups of genera and species wrapped by different combinations of their characteristics, however, are not necessarily harmonized with tribal or subtribal classifications available. Anatomical similarities and dissimilarities presented in this paper, as well as relationships among taxa presented in the classifications available, will be critically evaluated in the light of results of ongoing molecular phylogenetic analyses. Electronic Publication     

Pollen morphology of the Acalyphoideae and Euphorbioideae (Euphorbiaceae) of the Caatinga ecoregion in Brazil

This study dealt with six species of Acalyphoideae and 18 species of Euphorbioideae occurring in the Caatinga ecoregion, with emphasis on endemic species. Pollen samples were obtained from herbarium specimens and were acetolysed and analysed via light and scanning electron microscopy. The pollen of three genera of Acalyphoideae was medium to large, 3-colporate or 3-colpate, with an echinate-perforate exine that was reticulate, bireticulate, and microreticulate. The six genera of Euphorbioideae studied exhibited pollen grains that were small, medium and large; 3-colporate with margines; and an exine with microreticulate, microreticulate-rugulate, microreticulate-caveate, and reticulate ornamentation. The pollen characteristics were more variable in the Acalyphoideae compared to the relatively homogeneous Euphorbioideae. This study provides new data and interpretations of the pollen morphology of two subfamilies of the Euphorbiaceae of the Caatinga ecoregion in Brazil.     

Chemotaxonomy of the tribe Antidesmeae (Euphorbiaceae): antidesmone and related compounds

Selected species of the tribe Antidesmeae (Euphorbiaceae, subfamily Phyllanthoideae) have been screened for antidesmone occurrence and its content by quantitative HPLC (UV) and qualitative LC-MS/MS analysis. The LC-MS analysis allowing the additional detection of 17,18-bis-nor-antidesmone, 18-nor-antidesmone, 8-dihydroantidesmone and 8-deoxoantidesmone was carried out in the selected reaction monitoring (SRM) mode. Leaf material from herbarium specimens of 13 Antidesma spp., Hyeronima alchorneoides and Thecacoris stenopetala (all subtribe Antidesminae), as well as Maesobotrya barteri, Aporosa octandra (both Scepinae) and Uapaca robynsii (Uapacinae) were analysed. Additionally, freshly collected samples of different plant parts of two Antidesma spp. were investigated to ensure the significance of the results on herbarium specimens and to compare the antidesmone content in bark, root and leaves. Antidesmone could be unambiguously identified in 12 of 13 Antidesma spp., as well as in the two other investigated genera of subtribe Antidesminae, in levels of up to 65 mg/kg plant dry weight. Antidesmone was not found in specimens from other subtribes. Antidesmone-derived compounds occur in much lower concentrations than antidesmone.     

Molecular phylogenetics of tribe Poranthereae (Phyllanthaceae; Euphorbiaceae sensu lato)

Novel insights into the evolutionary history of a taxonomically complex tropical plant group were gained in this study using DNA sequence data. A molecular phylogenetic analysis of the newly circumscribed and expanded tribe Poranthereae (Phyllanthaceae) is presented. Sampling included 97 accessions for 63 of c. 120 species. Largely congruent results have been obtained from nuclear ribosomal ITS and plastid matK sequences. These analyses support the recognition of Andrachne and Leptopus as distinct genera. The deceptively similar Andrachne section Phyllanthopsis, Andrachne ovalis, and Leptopus decaisnei are separate lineages to be segregated. Zimmermannia and Zimmermanniopsis are embedded in Meineckia; Oreoporanthera is embedded in Poranthera; and Archileptopus is embedded in Leptopus. Andrachne section Pseudophyllanthus is polyphyletic, the two Madagascan endemics emerging as a sister clade to Meineckia. The noncontiguous distributions of Andrachne sensu lato and Leptopus sensu lato were found to be the result of separate evolutionary histories of morphologically similar clades, whereas Andrachne sensu stricto and Meineckia remain geographically disjunct. Actephila and Leptopus are sisters with a sympatric distribution in humid Asia. Presence of petals appears to be plesiomorphic for the tribe. Petals are reduced in Actephila and Oreoporanthera and lost in the Meineckia clade.     

Synopsis of the tribe Bocageeae (Annonaceae), with revisions of Cardiopetalum,Froesiodendron, Trigynaea,Bocagea, and Hornschuchia

The tribal name Bocageeae Endlicher is reestablished and the tribe is circumscribed on the basis of solitary internodal ebracteate pedicels that are articulated at the base, and pollen shed in polyads of eight or more grains. Septate anther locules, large pollen size, and seed appendages are prevalent in the tribe. Intectate pollen with free-standing columellae, rare in Annonaceae, occurs not only in the genusTrigynaea but also in some species ofBocagea andHornschuchia. As defined here, the Bocageeae include seven neotropical genera:Cymbopetalum, Porcelia, Bocagea, Cardiopetalum, Froesiodendron, Hornschuchia, andTrigynaea. The latter five genera are revised and the treatments include ten new species:Cardiopetalum plicatum, Froesiodendron urceocalyx, Hornschuchia lianarum, H. santosii, H. leptandra, Trigynaea cinnamomea, T. lanceipetala, T. triplinervis, T. lagaropoda, andT. axilliflora, all from tropical South America. A new combination,Froesiodendron longicuspe, changes the rank of that taxon from subspecies to species.Cardiopetalum surinamense is removed fromFroesiodendron and reassigned toCardiopetalum on the basis of its connate petals, dehiscent monocarps, and seeds with bilobed arils. A cladogram provides an explicit hypothesis of intergeneric relationships in the tribe. The new combinationOnychopetalum periquino, based onTrigynaea periquino, is made.     

Glyphostylus (Euphorbiaceae, Hippomaneae) reexamined

The monotypic genus Glyphostylus (Euphorbiaceae) is compared with Excoecaria , and it is concluded that both genera are not distinct. Glyphustylus , instead, exhibits all characters diagnostical for Excoecaria. Gl.vphostylus laoticus is considered as a species of Excoeearia . and the necessary combination is made.     

Synopsis of Boraginaceae subfam. Boraginoideae tribe Boragineae in Italy

A synopsis of the Italian taxa of tribe Boragineae (Boraginaceae, subfam. Boraginoideae) is given as a second contribution to the treatment of the family for the Flora Critica d'Italia project. The work is mainly based on the critical study of herbarium material and extensive literature survey. All relevant floristic reports were examined and types of all the accepted (51) and most (37/46) of the synonymized names of taxa, specific and infraspecific, reported from the National territory are indicated. In the light of karyological and morphological evidence, the new combination Pulmonaria vallarsae subsp. apennina is proposed. As a result, 12 genera and 37 species are recognized, of which 7 are allochtonous and 6, plus two subspecies, are endemic. A synthetic floristic treatment is provided, including analytical keys, hybrids, list of synonyms and short distribution notes. In addition, detailed distribution maps are provided, together with the lists of the selected specimens upon which they are based.     

Pollen morphology of the Flueggeinae (Euphorbiaceae,Phyllanthoideae)

A total of 129 species from the subtribe Flueggeinae of the tribe Phyllantheae (Euphorbiaceae, Phyllanthoideae) were investigated using light and scanning electron microscopy, and 10 species using transmission electron microscopy, in order to evaluate the relationships between the eight constituent genera: Breynia, Flueggea, Glochidion, Margaritaria, Phyllanthus, Reverchonia, Richeriella, and Sauropus. Of these genera, Flueggea, Margaritaria and Richeriella share pollen with a prolate spheroidal meridional outline and a 3-colporate aperture system. Pollen of Reverchonia is also 3-colporate, but differs from that of the Flueggea alliance by its clearly prolate shape, tilioid ornamentation and absence of costae endopori. Breynia and Sauropus have 4–12 and 3–16-colporate pollen, respectively, with diploporate colpi. Two pollen types are recognised in Breynia, and four in Sauropus, one of which supports the recognition of Sect. Hemisauropus. Glochidion pollen is 3–6-colporate, and similar to that of Breynia in having reticulate sculpture with Y-shaped sexine structures, but it has monoporate colpi. Of the genus Phyllanthus, only species with pollen with diploporate colpi have been studied. Seven types are described. Diploporate Phyllanthus pollen can be distinguished from that of Breynia and Sauropus by its distinct colpus margins consisting of parallel muri. Colpal irregularities and endoaperture configurations in the subtribe are discussed, and pollen morphological trends are hypothesised. Placed in the successiform aperture series, the Flueggea alliance and Reverchonia form a basal group. Glochidion is considered intermediate, giving rise to the Breynia-Sauropus group. The relationship with Phyllanthus remains unclear.     

A new tribe, Epistomentini, of Buprestidae (Coleoptera) with a redefinition of the tribe Chrysochroini

A new tribe Epistomentini is designated to include Epistomentis Solier, Cyria Solier (with which Cyrioides Carter and Xenocyria Obenberger are synonymized), Diadoxus Thomson, Cyrioxus Hoschek, and a new genus and species Araucariana queenslandica, which is described. The tribe Chrysochroini is redefined, and Epidelus Deyrolle, Cyalithus Thomson, Cyalithoides Fisher and Chrysopistus Thery are transferred to it from the Chalcophorini. Chalcophoropsis Thomson is transferred from the Chrysochroini to the Chalcophorini. A reassessment of the value of some characters used in the higher classification of the Buprestidae is given.     

Molecular phylogenetics of the giant genus Croton and tribe Crotoneae (Euphorbiaceae sensu stricto) using ITS and TRNL-TRNF DNA sequence data

Parsimony, likelihood, and Bayesian analyses of nuclear ITS and plastid trnL-F DNA sequence data are presented for the giant genus Croton (Euphorbiaceae s.s.) and related taxa. Sampling comprises 88 taxa, including 78 of the estimated 1223 species and 29 of the 40 sections previously recognized of Croton. It also includes the satellite genus Moacroton and genera formerly placed in tribe Crotoneae. Croton and all sampled segregate genera form a monophyletic group sister to Brasiliocroton, with the exception of Croton sect. Astraea, which is reinstated to the genus Astraea. A small clade including Moacroton, Croton alabamensis, and C. olivaceus is sister to all other Croton species sampled. The remaining Croton species fall into three major clades. One of these is entirely New World, corresponding to sections Cyclostigma, Cascarilla, and Velamea sensu Webster. The second is entirely Old World and is sister to a third, also entirely New World clade, which is composed of at least 13 of Webster's sections of Croton. This study establishes a phylogenetic framework for future studies in the hyper-diverse genus Croton, indicates a New World origin for the genus, and will soon be used to evaluate wood anatomical, cytological, and morphological data in the Crotoneae tribe.     

Taxonomic revision of the tribe Hymenodictyeae (Rubiaceae, Cinchonoideae)

The palaeotropical tribe Hymenodictyeae Razafim. & B. Bremer, belonging to the otherwise predominantly Neotropical subfamily Cinchonoideae  s.s. (coffee family or Rubiaceae), is revised here. The tribe as presently circumscribed contains two genera, Hymenodictyon Wall. and Paracorynanthe Capuron ex J.-F. Leroy, and is distinct from the other Cinchonoideae tribes with capsular fruits in having stipules bearing large, deciduous colleters on the margins, valvate corolla aestivation, and lenticellate capsular fruits that contain elongate, bilaterally flattened, and accrescent placentae. We recognize 22 Hymenodictyon species, including four new species ( H. antakaranensis sp. nov. , H. epiphyticum sp. nov. , H. madagascaricum sp. nov. , and H. tsingy sp. nov. ), and the two described species of Paracorynanthe . Hymenodictyon is distinguished from its sister genus, Paracorynanthe , by simple or compound spicate, racemose or thyrsoid inflorescences and corolla lobes without any appendages, rather than compound umbelliform inflorescences and corolla lobes prolonged by ciliate appendages. A full taxonomic treatment, keys, and distribution maps of all recognized Hymenodictyon and Paracorynanthe species are provided. Five lectotypes and one neotype are designated. Finally, six Hymenodictyon species are illustrated for the first time.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 331–386.     

The mis-measure of manioc (Manihot esculenta,Euphorbiaceae)

The distinction between “bitter” and “sweet” (toxic and non-toxic) varieties of the cyanide-containing food crop manioc (Manihot esculenta, Euphorbiaceae) has a long tradition in the tropical forest areas of South and Central America where it was first cultivated. Yet this distinction has no taxonomic basis. The levels of cyanogenic glucosides found in manioc varieties not only vary widely, but do not correspond with any other known morphological or ecological feature. Nonetheless, these two “varieties” are commonly reported to have distinct geographical and cultural distributions and each is associated with a particular traditional food complex. This paper reviews the literature regarding the nature, distribution, and traditional uses of manioc varieties and concludes that (1) the geographical and cultural distribution of bitter and sweet varieties of manioc may not be as distinct as has been thought; (2) traditional categories of bitter and sweet manioc may stem more from culturally derived belief systems than from actual known levels of toxicity; and (3) the choice of complex and labor intensive processing methods usually associated with bitter manioc may not be required for detoxification but rather for the derived food products, particularly manioc flour.     

Synopsis of Satanocrater (Acanthaceae)

Four species are recognized in the tropical African genus Satanocrater: S. fellatensis (= S. berhautii) in Sudan, Ethiopia and Guinea, S. ruspolii in Ethiopia, Somalia and Kenya, S. somalensis (= S. coccineus) in Ethiopia and Somalia, and S. paradoxus in Ethiopia, Somalia and Kenya. Three lectotypes and one neotype are selected. A key to the species and an illustration of S. ruspolii are provided.     

Leaf structure of species from three closely related genera from tribe Crotoneae Dumort. (Euphorbiaceae s.s., Malpighiales)

Euphorbiaceae s.s. is one of the largest angiosperm families, comprising nine major lineages. Molecular analysis indicated a polyphyletic condition in the subfamily Crotonoideae s.l., with four lineages identified, including one termed inaperturate crotonoids, which encompasses the tribe Crotoneae. Anatomical data were obtained from 15 species of Croton sect. Cyclostigma (Griseb.) Müll. Arg., Brasiliocroton P. Berry & I. Cordeiro and Astraea Klotzsch, which belongs to this lineage. The presence and protodermal origin of the idioblasts with lipophilic content and the interruption of the palisade by collenchyma layers on the adaxial side of the mid vein are common to the Croton L. species analysed and Brasiliocroton. The presence of dorsiventrality, paracytic stomata, collateral vascular bundles, branched non-articulated laticifers and crystal idioblasts in the species studied indicates morphological similarities among the three genera. The interpretation of the stipite of the complex trichomes as emergences is a structural novelty for the group. Leaf anatomy provided an excellent source of characters for future studies of the tribe Crotoneae (Euphorbiaceae s.s., Malpighiales).     

Synopsis of Adenanthera (Leguminosae-Mimosoideae)

The generic relationship, the palynology and the infrageneric classification of the tropical Asian-Australian-Melanesian genus Adenanthera is discussed. Twelve species are recognized. Adenanthera microsperma and A. tamarindifolia are reinstated. A. kostermansii and A. marina are described as new, and A. malayana is divided in two subspecies, ssp. malayana from Malaya and Sumatra and ssp. andersonii from Borneo. All species are illustrated.