Phylogenetic position and revised classification of Acacia s.l. (Fabaceae: Mimosoideae) in Africa,including new combinations in Vachellia and Senegalia

Previous phylogenetic studies have indicated that Acacia Miller s.l. is polyphyletic and in need of reclassification. A proposal to conserve the name Acacia for the larger Australian contingent of the genus (formerly subgenus Phyllodineae) resulted in the retypification of the genus with the Australian A. penninervis. However, Acacia s.l. comprises at least four additional distinct clades or genera, some still requiring formal taxonomic transfer of species. These include Vachellia (formerly subgenus Acacia), Senegalia (formerly subgenus Aculeiferum), Acaciella (formerly subgenus Aculeiferum section Filicinae) and Mariosousa (formerly the A. coulteri group). In light of this fragmentation of Acacia s.l., there is a need to assess relationships of the non‐Australian taxa. A molecular phylogenetic study of Acacia s.l and close relatives occurring in Africa was conducted using sequence data from matK/trnK, trnL‐trnF and psbA‐trnH with the aim of determining the placement of the African species in the new generic system. The results reinforce the inevitability of recognizing segregate genera for Acacia s.l. and new combinations for the African species in Senegalia and Vachellia are formalized. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 500–523.     

Pollen morphology and phylogenetic relationships in neotropical Phyllanthus (Euphorbiaceae)

In order to provide new insights into phylogenetic relationships among the neotropical taxa of Phyllanthus , 28 illustrations are provided of the pollen grains of 22 selected species studied from 11 sections of the subgenera represented in the neotropics. Special attention has been given to subgenus Conami because of its variability in pollen morphology: of eight species illustrated, the apertures are diploporate colpi in three species and pores in five species; exine ornamentation is vermiculate in two species and pilate in the other six species. The six species in the neotropical sections Pityrocladus and Microglochidion (subgenus Emblica ) are characterized by prolate grains with an increased number of colpi (4–8). Of particular interest are species in which the pollen exine is clypeate (with exine shields); clypeate pollen grains are illustrated in two species of subgenus Xylophylla and in one species of section Cyclanthera that has unique exine shields with single central pila. The pollen of the one Brazilian phylloclade-bearing species illustrated (in section Choretropsis ) has 3-colporate grains with reticulate exine, typical for subgenus Phyllanthus , and very different from the clypeate grains of the West Indian phylloclade-bearing species in section Xylophylla . This pollen evidence clearly demonstrates homoplasy in the origin of phylloclades in Phyllanthus . Pollen morphological data suggest that the neotropical taxa of Phyllanthus have arisen following colonization from Africa (subgenus Kirganelia ) and Asia (subgenus Emblica ). © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 325–338.     

Molecular phylogenetics of Acacia (Fabaceae: Mimosoideae) based on the chloroplast MATK coding sequence and flanking TRNK intron spacer regions

The tribe Acacieae (Fabaceae: Mimosoideae) contains two genera, the monotypic African Faidherbia and the pantropical Acacia, which comprise about 1200 species with over 950 confined to Australia. As currently recognized, the genus Acacia is subdivided into three subgenera: subg. Acacia, subg. Aculeiferum, and the predominantly Australian subg. Phyllodineae. Morphological studies have suggested the tribe Acacieae and genus Acacia are artificial and have a close affinity to the tribe Ingeae. Based on available data there is no consensus on whether Acacia should be subdivided. Sequence analysis of the chloroplast trnK intron, including the matK coding region and flanking noncoding regions, indicate that neither the tribe Acacieae nor the genus Acacia are monophyletic. Two subgenera are monophyletic; section Filicinae of subgenus Aculeiferum does not group with taxa of the subgenus. Section Filicinae, eight Ingeae genera, and Faidherbia form a weakly supported paraphyletic grade with respect to subg. Phyllodineae. Acacia subg. Aculeiferum (s. s.) is sister to the grade. These data suggest that characters currently used to differentiate taxa at the tribal, generic, and subgeneric levels are polymorphic and homoplasious in cladistic analyses.     

Two new species of Acacia (Leguminosae–Mimosoideae) from Ethiopia and Yemen

Two new species of Acacia belonging to subgenus Acacia, A. origena from the high lands of N Ethiopia and Yemen and A. prasinata from the Rift Valley in central Ethiopia are described. The external morphology of the latter species is shortly described.     

Phylogeny of the Afro-Madagascan Aristea (Iridaceae) revisited in the light of new data on pollen morphology

Scanning and electron microscopic studies of the pollen grains of several species of the sub-Saharan African and Madagascan genus Aristea of c . 50 species, supplement data from an earlier study and enable us to include these species in a phylogenetic analysis. TEM examination of a range of pollen grains of representative pollen types in Aristea makes it possible to reinterpret past SEM results so that apertures covered with exine masses and previously called sulculate, are now seen to be either zonasulculate or dizonasulculate. Revised and expanded data are combined in a matrix together with new data for seed morphology in the genus, which like the capsules is remarkably diverse, including angular seeds with reticulate sculpturing, lamellate or triangular-columnar seeds with smooth or reticulate surfaces and with smooth or papillate or areolate margins. This new phylogenetic analysis suggests that Aristea should remain treated as comprising three subgenera. In subgenus Eucapsulares we add a new section Latifolieae for A. latifolia , which seems an isolated relic restricted to the mountains of the Cape Region of South Africa, an area of winter rainfall. Species with similar unspecialized capsule and seed morphology occur only in eastern south and tropical Africa and Madagascar, but they have derived pollen morphology whereas the pollen of A. latifolia is plesiomorphic. This species appears to be the closest living ancestor of the genus. In subgenus Aristea we continue to recognize three sections, sect. Racemosae , sect. Singulares and sect. Aristea. Nesting of subgenus Pseudaristea within section Racemosae of subgenus Aristea is probably an artefact of the analysis and does not receive bootstrap support, and hence we continue to recognize this group of species at subgeneric rank.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 41–68.     

Appomattoxia ancistrophora gen. et sp. nov., a new Early Cretaceous plant with similarities to Circaeaster and extant Magnoliidae

A new genus and species of fossil angiosperm (Appomattoxia ancistrophora) is established based on well-preserved fruiting units and associated pollen from the Early Cretaceous (Early or Middle Albian) Puddledock locality in the Potomac Group sequence of Virginia, eastern North America. Fruiting units are small, unilocular, and with a single, pendulous, orthotropous seed. The fruit surface is characterized by densely spaced unicellular spines with hooklike tips, which probably functioned in biotic dispersal. Pollen grains adhering to the stigmatic area of many specimens are monocolpate and tectate with granular to columellate infratectal structure, and are similar to dispersed grains assigned to Tucanopollis and Transitoripollis. Comparison of fossil Appomattoxia ancistrophora with extant plants reveals an unusual combination of characters that includes similarities with some magnoliid taxa, particularly Piperales (Piperaceae, Saururaceae) and Laurales (Chloranthaceae), as well as the monotypic ranunculid family Circaeasteraceae. Appomattoxia ancistrophora differs from extant Piperales in having a pendulous rather than erect ovule, and differs from extant Circaeaster in details of the fruit wall, as well as the presence of monosulcate rather than tricolpate pollen.     

Erdtmanitheca portucalensis,a new pollen organ from the Early Cretaceous (Aptian–Albian) of Portugal with Eucommiidites-type pollen

A new lignitised, slightly compressed pollen organ, Erdtmanitheca portucalensis, with affinities to extinct Erdtmanithecales from the Early Cretaceous (Aptian–Albian) of Vale de Água (Lusitanian Basin, western Portugal), is described. The pollen organ is composed of loosely arranged microsporophylls radiating from a central core. The estimated number of microsporophylls is about 100–150. The microsporophylls are sessile and ellipsoidal to barrel-shaped with a flattened or slightly apically depression containing about ten narrow sporangia. The sporangia enclose abundant well-preserved pollen grains of Eucommiidites-type. Pollen grains found in situ are elliptical in equatorial outline, about 16.0–27.2 μm long and 11.9–16.4 μm wide. The main (distal) colpus is long with expanded rounded ends. It is flanked by two subsidiary colpi in an almost equatorial position. The surface of the pollen wall is psilate and occasionally punctate. The ektexine is composed of a distinct tectum, granular infratectal layer and a thin foot layer. The endexine is thick and laminar. The new Early Cretaceous Portuguese pollen-organ is similar in several respects to that of Erdtmanitheca texensis described from the Late Cretaceous of Texas, USA. The new fossil species further documents the importance of the Bennettitales-Erdtmanithecales-Gnetales group in the Early Cretaceous floras of Portugal extending the stratigraphic and geographical distribution of the genus with regard to systematic and phylogenetic significance of the Eucommiidites-producing plants that may have been co-occurring with the Early Cretaceous diversification of angiosperms. It is ascertained that perforate tectum occurs in pollen grains with a well-developed foot layer as well as in pollen grains in which a foot layer is poorly developed or lacking, and that pollen features do not support a separation of the Erdtmanithecales seeds and pollen organs.     

Pollen morphology of the Phyllanthus species occurring in the continental United States

Twelve native United States species have been examined. The pollen grains could be divided into five pollen types based on sexine ornamentation, aperture structure, sexine/nexine ratio and sexine 1/sexine 2 ratio. Within these types it was often possible to distinguish the grains of the different species. Two groups of types have been recognized based on the similarities in ornamentation. One group, consisting of the Phyllanthus amarus type, the P. tenellus type and the P. urinaria type, is characterized by the Tilia structure, whereas the second group of types, comprising the P. caroliniensis type and the P. liebmannianus type are microreticulate. The postulation that different subgenera show different pollen types is true in general, but there is one exception: P. niruri, taxonomically belonging to the subgenus Phyllanthus, has pollen grains of the P. caroliniensis type, a type also comprising species from the subgenus Isoclades.     

Cotyledon architecture and anatomy in the Acacieae (Leguminosae: Mimosoideae)

Cotyledon size, shape, venation pattern and anatomy have been investigated in Faidherbia albida and 152 species of Acacia representing the three subgenera Acacia, Aculeiferum and Heterophyllum. Cell volumes of epidermis, palisade and storage tissue, stomatal frequency, stomatal index and frequency of stomatal types have been determined for F. albida and 12 species from each subgenus. The data obtained support the recognition of the subgenera of Acacia as separate taxa but provide no indication of their status. The evidence from cotyledons also supports the separation of Faidherbia from Acacia , and the amalgamation of the Acacieae and Ingeae.     

Callionymus omanensis,a new species of dragonet from Oman,north‐western Indian Ocean (Teleostei: Callionymidae)

A new species of deep‐living dragonet Callionymus omanensis from Oman is described on the basis of a single male specimen collected in a trawl from 500 m depth off the coast of Oman. The new species is characterized within the subgenus Bathycallionymus by having a small branchial opening; head short (3·9 in proportion to standard length); eye large (2·4 in proportion to head length); preopercular spine with a long, upcurved main tip, with a small antrorse barb and a larger antrorse spine, and with a strong antrorse spine laterally at the preopercular‐spine base, ventral margin smooth; first dorsal fin slightly higher than second dorsal fin (male); second dorsal fin distally straight; 17 pectoral fin rays; distal end of caudal fin slightly pointed, with two median unbranched rays bearing short filaments; first dorsal fin with basal black spot reaching from first to fourth membranes, third membrane with an ocellated distal black blotch; second dorsal fin with vertical dark grey bars; distal three‐fourths of anal fin black; upper half of caudal fin with oblique dark grey bars; pelvic fin dark grey, second ray basally with a black blotch. The new species is compared with similar species. Revised keys to callionymid species of the western Indian Ocean and the Red Sea, as well as species of the subgenus Bathycallionymus, are presented.     

FOSSIL FLORAL EVIDENCE OF MALPIGHIACEAE AND AN EARLY PLANT-POLLINATOR RELATIONSHIP

Fossil flowers with affinities to Malpighiaceae have been discovered in the Middle Eocene Claiborne formation of northwestern Tennessee. The new taxon Eoglandulosa warmanensis gen. et sp. nov. Taylor and Crepet, has paired glands on the five sepals, clawed petals and tricolporate pollen with reticulate ornamentation and an unusual infratectal wall structure of anastomosing elements. The fossil is similar in wall structure to some extant species of Malpighiaceae. Glandular floral morphology in extant species is associated with specific anthophorid bee pollinators and the fossil evidence suggests that such specific plant-pollinator relationships existed during the Eocene. This fossil species also suggests that by the Eocene, South American floral elements had migrated to North America via island pathways, and that the Mississippi embayment was nearly frost-free.     

四种番荔枝科植物花粉形态

利用扫描和透射电子显微镜,观察了番荔枝科(Annonaceae)4属4种植物的花粉形态与结构.刺果番荔枝(Annona muricata)和金钩花(Pseuduvaria trimera)为四合花粉,有四角形、偏菱形、T-型、十字形和四面体形.前者花粉表面具小穿孔,后者为皱波状纹饰.蕉木(Oncodostigma hainanense)和那大紫玉盘(Uvaria macclurei)为单花粉,前者表面为皱波状纹饰,后者为粗褶皱状纹饰.透射电子显微镜下,蕉木和那大紫玉盘覆盖层较厚、连续.蕉木覆盖下层较薄,偶尔可见颗粒状物质或不规则小柱,为过渡型覆盖下层;那大紫玉盘覆盖下层颗粒状.基层分为内外两层,外层较厚,平或呈波浪状,内层具有2～4片层结构.在不同属或同属不同种之间,花粉特征差异较大,多样性非常丰富.金钩花具有四合花粉、覆盖下层小柱状等进化特征,同时又具有花粉粒小、覆盖层无穿孔等较原始特征.     

Past and present vegetation ecology of Laetoli, Tanzania

We are attempting to set up a new protocol for palaeoecological reconstruction in relation to the fossil hominin site Laetoli, Tanzania. This is based on the premise that habitat variability in the past was at least as great as at present; that this variability at the landscape level is a function of variations in geology, soils, and topography rather than climate; and that vegetation type at the landscape level can be reconstructed from these environmental variables. Measurable variation in climate in tropical Africa today occurs over distances of at least 100 km, so that ranges of habitat variation within the limited area of Laetoli today can be reconstructed in relation to soils and topography, and the effects of climate changes are then estimated in relation to these other factors. In order to document the modern vegetation, we have made voucher collections of plants in the Laetoli region, recorded distributions of plants by habitat, climate, soil, and topography, and mapped the vegetation distributions. Results show that areas of low relief have soils with impeded drainage and dense Acacia drepanolobium woodland, having low canopies when disturbed by human action, higher when not; shallow brown soils on volcanic lavas have four woodland associations, two dominated by Acacia species, two by Combretum-Albizia species; shallow volcanic soils to the east have a woodland association with Croton-Dombeya-Albizia species; elevated land to the east on volcanic soils has two associations of montane-edge species, one with Croton-Celtis-Lepidotrichilia, and the other with Acacia lahai; the eastern highlands above 2,750 m have montane forest; seasonal water channels flowing from east to west have three Acacia riverine woodland associations; three deep valleys to the north of the area have dense riverine woodland with Celtis, Albizia, Euclea, Combretum, Acacia spp.; emergence of springs at Endulen feed a perennial stream with closed gallery forest with Ficus-Croton-Lepidotrichilia; and, finally, recent ash falls have produced immature alkaline soils with calcrete formation and short grass vegetation. All of these vegetation associations have been modified by human disturbance to greater or lesser degrees, and we have attempted to allow for this both by basing the associations on the least modified areas and by predicting how the associations, or parts of associations, have been altered by human action. Past land forms at Laetoli have been based on the geology and geomorphology of the area. Past vegetation patterns were estimated by superimposing present distributions of plant associations on equivalent landforms in the past, assuming similar climate to the present. This indicates the overall pattern of vegetation at Laetoli to have been a mosaic of low and tall deciduous woodlands and with riverine woodland and forest associations along water courses. Low woodlands would have been dominated by Acacia species, and tall woodlands by Combretum-Albizia species, with increasing increments of montane species, such as Croton species, to the east of the area. Riverine woodlands would have been dominated by Acacia-Euclea species, with wetter associations (downriver or linked with spring activity) supporting gallery forest with Ficus, Celtis, and Croton species. These are all species associations common in the area today, and with landforms little changed in the past, and assuming similar climate, there is every reason to predict that they would have been present in the past. Moreover, Pliocene environments lack the human disturbance that has destroyed much of the present day vegetation. Presence of woodlands is supported by fossil wood attributed to several of the tree species present in the area today and by similarities in the mammalian community structure between past and present. Having established the pattern for Pliocene vegetation based on climatic variables existing today, we then predict the effects of past variations in climate.     

Phylogeny of the subgenus Culicoides and related species in Italy, inferred from internal transcribed spacer 2 ribosomal DNA sequences

Biting midges of the genus Culicoides (Diptera: Ceratopogonidae) include vectors for the economically important animal diseases, bluetongue (BT) and African horse sickness (AHS). In the Mediterranean Basin, these diseases are transmitted by four species of Culicoides: the first three belong in the subgenus Avaritia Fox and are Culicoides imicola Kieffer, Culicoides obsoletus (Meigen) and Culicoides scoticus Downes and Kettle; the fourth is Culicoides pulicaris (Linnaeus) in the subgenus Culicoides Latreille. In the Palaearctic Region, this subgenus (usually referred to as the C. pulicaris group) now includes a loose miscellany of some 50 taxa. The lack of clarity surrounding its taxonomy stimulated the present morphological and molecular study of 11 species collected in Italy. Phylogenetic analysis of nuclear ribosomal DNA internal transcribed spacer 2 (ITS2) sequence variation demonstrated a high degree of divergence. These results, combined with those from a parallel morphological study, disclosed: (1) that some previously described taxa should be resurrected from synonymy; (2) that there are new species to be described; (3) that the subgenus Culicoides (as currently employed) is a polyphyletic assemblage of four lineages - the subgenus Culicoides sensu stricto, the subgenus Silvicola Mirzaeva and Isaev, the subgenus Hoffmania Fox and the hitherto unrecognized Fagineus species complex. Each is discussed briefly (but not defined) and its constituent Palaearctic taxa listed. Strong congruence between morphological and molecular data holds promise for resolving many of the difficult taxonomic issues plaguing the accurate identification of vector Culicoides around the world.     

Cyanogenesis in Acacia pachyphloia

The cyanogenic glycoside, proacacipetalin, is reported from Acacia pachyphloia (Acacia subgenus Acacia). This represents the first record of a glycoside with an aliphatic aglycone from a species of Acacia indigenous to Australia. This finding reinforces the taxonomic distinctions between subgenus Acacia and subgenus Phyllodineae.     

Pollen structure, tetrad cohesion and pollen-connecting threads in Pseuduvaria (Annonaceae)

The structure of the pollen of 42 species of Pseuduvaria (Annonaceae) is described. The pollen is consistently inaperturate, isopolar and radially symmetrical. Four basic patterns of exine sculpturing are identified: rugulate, verrucate, scabrate and psilate. The exine stratification of one representative species, P. macrocarpa , is shown to be entirely ectexinal. The ectexine consists of a discontinuous outer tectal layer, a columellar infratectal layer, and an inner lamellar foliated foot layer; the intine is very thin and fibrillar. The pollen is invariably released as acalymmate tetrads, in which the tectum is absent from the proximal walls. The individual pollen grains within the tetrads are connected by crosswall cohesion, involving both exine and intine; this form of cohesion has not hitherto been reported in the Annonaceae. In addition, pollen grains of neighbouring tetrads are connected in two different ways, viz. short exine connections and non-sporopollenin pollen-connecting threads. Neither of these cohesion mechanisms has previously been reported for the genus. The function of the various forms of cohesion between pollen grains and tetrads in Pseuduvaria is discussed as a mechanism to enhance the efficiency of pollination by enabling the fertilization of multiple ovules following a single pollinator visit.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 69−78.     

Two leguminous forest trees new to the flora of French Guiana

Stryphnodendron moricolor (Mimosaceae) is described, illustrated and discussed. The genusMonopteryx (Fabaceae) is reported for the first time outside Amazonian Brazil and adjacent Venezuela. A key distinguishing the three known species ofMonopteryx is presented.     

Domicile constructing phlaeothripine Thysanoptera from Acacia phyllodes in Australia: Dunatothrips Moulton and Sartrithrips gen.n., with a key to associated genera

Abstract. Domiciles are constructed by adult females of species in several genera of Australian phlaeothripine Thysanoptera. This is done by glueing or sewing together two or more Acacia phyllodes, or by weaving a tentlike structure on a phyllode surface. These thrips then breed within the domiciles, sometimes for multiple generations. Mature females become dealate in seven of twelve species discussed, through fracture of the distal four-fifths of each wing, although young females and all males have complete wings. Of these twelve species, six are placed in Dunatothrips Moulton and six in a new genus, Sartrithrips . Adults of the latter genus have a spinelike structure dorsal to the anus, 'the supra-anal process', that is not found in any other member of Phlaeothripidae. Despite the many structural and biological character states shared by these twelve species, molecular methods using one nuclear gene ( wingless ) and one mitochondrial gene (cytochromome oxidase 1) demonstrated that the two genera are monophyletic but do not have a sister-group relationship. Identification keys are provided to nineteen described phlaeothripine genera that are known to be associated with Acacia phyllodes, and to the twelve species in Dunatothrips and Sartrithrips , of which ten are described as new.     

Pollen morphology and structure of Tropical and Subtropical American genera of the Piptadenia -group (Leguminosae: Mimosoideae)

A survey of the pollen morphology of the American taxa of the Piptadenia -group (Mimoseae) was done including all together 384 species of the following genera: Piptadenia , Parapiptadenia , Pseudopiptadenia , Piptadeniopsis , Microlobius (= Goldmania ), Stryphnodendron , Adenopodia , Anadenanthera and Mimosa . Because of a high morphological and structural variation the definition and characterization of pollen types turned out to be very difficult. Nevertheless the presence and durability of some pollen characters allow a provisional grouping: (1) always tetrads or polyads, (2) small size 6 (20) 40 mum, (3) unstable shape but currently ovoidal and, if biconvex, ellipsoidal in the outline, (4) number of pollen grains of the polyads very variable and unstable; the most frequent numbers are 8-12-16, (5) pollen grains irregularly united in polyads, with homomorphous exine and, (6) pores always located in the angles of the pollen grains in distal-subdistal position, rarely subdistal, (7) without or with pseudocolpi or subpseudocolpi (never colpi) in the distal face, (8) endexine forming an annulus around the endoaperture with ends of free lamellae with a central white line, (9) the tectal stratum above the pores forming a small vestibular space (except in Anadenanthera ). These palynological characters or most of them confirm the affinity among genera in the Piptadenia -group and its distinction from other taxa of the subfamily Misosoideae with tetrads or polyads.