A survey of species assigned to the fossil pollen genus Anacolosidites

Anacolosidites Cookson & Pike, a fossil pollen genus recorded since the Campanian, is peculiar in its morphology – six‐porate with three apertures on each hemisphere, located away from the equator, and with the distal and proximal apertures positioned over each other. Representatives of this fossil genus are widely considered to represent extant Olacaceae from tribe Anacoloseae. Olacaceae is an exclusively tropical angiosperm family with a pantropical distribution; consequently the fossils are often used to suggest a tropical climate and in addition are frequently used as a stratigraphic marker. Fossil species assigned to Anacolosidites are quite variable and may not all represent Olacaceae, in which case they may not indicate tropical climate.

The present study is a morphological survey of fossil pollen assigned to Anacolosidites; it identifies the published reports of the fossil species that probably represent positive occurrences of Olacaceae pollen in the fossil record. Within Olacaceae, Anacolosidites‐type pollen is usually compared with pollen of genera in tribe Anacoloseae, in particular: Cathedra, Anacolosa and Phanerodiscus, but never with the pollen of Ptychopetalum, a genus from tribe Olaceae with closely similar pollen to the other three genera, but with a reticulate tectum and very small circular apertures located near the equator. Nevertheless, the records of reticulate Anacolosidites species which have been excluded from the emended diagnosis are unlikely to be related to Ptychopetalum. The earliest accepted record of Anacolosidites is from the Late Cretaceous (Maastrichtian) of Germany. However, most Late Cretaceous records, and later Russian and Chinese occurrences referred to Anacolosidites, consist mainly of Normapolles‐type pollen, whereas many of the Cenozoic records assigned to Anacolosidites have a much clearer affinity with the pollen of Anacolosa, Cathedra and Phanerodiscus (tribe Anacoloseae). The newly emended genus Anacolosidites may be used as a stratigraphic marker for tropical or megathermal climatic conditions.     

Pollen morphology of the Pavetteae (Rubiaceae,Ixoroideae) and its taxonomic significance

Pollen grains of the tribe Pavetteae (Rubiaceae, subfamily Ixoroideae) are examined using LM and SEM. Grains are 3‐ or 4‐colporate and (semi‐) tectate (in one Versteegia species atectate). Sexine patterns vary between perforate, microreticulate, reticulate, rugulate and striato‐reticulate. Supratectal elements are sometimes present. The variation in pollen morphology in the Pavetteae allows to recognize seven pollen types, the distribution of which is useful to evaluate generic delimitations and relationships within the tribe. Pollen characters corroborate the close relationships between the genera Coleactina, Dictyandra and Leptactina and between Homollea, Homolliella and Paracephaelis. All the genera of the tribe proved to be stenopalynous (the species examined possess the same pollen type), except Pavetta, Rutidea, Versteegia and Tarenna which are eurypalynous. In the huge genus Pavetta the existing infrageneric classification is supported pollen morphologically. Pollen morphology further indicates that the genus Tarenna is badly delimited and strongly in need of a revision. The small genus Versteegia is in need of further taxonomic and palynological study to understand the pollen morphological variation encountered here. At a higher rank, pollen morphology also does not contradict the recent division of the Pavetteae in the Ixoreae (a stenopalynous tribe with presumably primitive pollen) and the Pavetteae sensu stricto (eurypalynous).     

Pollen morphology and systematics of Burseraceae

The Burseraceae are a medium‐sized family in which 18 genera are currently recognised. They are the subject of a long‐term project to describe the pollen morphology from light, scanning electron and transmission electron microscopy. The pollen morphology of tribe Protieae has been published, as well as an account of the pollen of the African taxa in the family. Pollen data for the other two tribes, Bursereae and Canarieae, are more or less complete. The pollen of all the genera have been examined, with the exception of the recently described Pseudodacryodes Pierlot for which, currently, there is no pollen material available. This paper summarises the results.

There is considerable variation in exine and aperture features between, and occasionally within, the genera and 14 major pollen types are defined, including two previously undescribed types: ‘Canarium oleiferum’ and ‘Canarium gracile’. The distribution of pollen characteristics throughout the family is compared with previously published tribal and subtribal groupings, as well as with current ideas of generic relationships from molecular analyses. Comparisons show notable congruence of pollen data with molecular data. To some extent pollen morphology is different for each of the subtribes. Nevertheless, there are some notable exceptions, for example, the pollen of Garuga and Boswellia are remarkably similar, although Garuga has been included, somewhat tenuously, in tribe Protieae, and Boswellia is included in tribe Bursereae, subtribe Boswelliinae. In a recent molecular tree Garuga and Boswellia appear to be closely related, and this supports the conclusion, based on several macromorphological characters as well as pollen, that Garuga should be transferred to tribe Bursereae.     

POLLEN MORPHOLOGY OF THE TRIBE SWARTZIEAE (SUBFAMILY PAPILIONOIDEAE: LEGUMINOSAE). 1. INTRODUCTION AND ALL GENERA EXCLUDING ALDINA AND SWARTZIA

The pollen morphology of 9 of the 11 genera of the tribe Swartzieae is described together with that of Holocalyx and Cyathostegia, two genera recently removed from the tribe based on macromorphology. The pollen is small, spheroidal to subprolate, primarily tricolporate with a perforate tectum and generally typical of the Leguminosae. Nonetheless, many of the genera have distinctive pollen morphology. Baphiopsis is 6-colporate. The genera Harleyodendron, Lecointea and one species of Exostyles have supratectal spinules. Bocoa viridiflora has striate/rugulate ornamentation very distinct from the other species of the genus Bocoa. The exine stratification is varied but Candolleodendron has a very thick endexine and narrow foot layer. The pollen of African taxa does not differ significantly from that of South American taxa. Pollen morphology does not clarify the taxonomic relationships of the tribe and provides little evidence to assist in positioning Holocalyx and Cyathostegia.     

Pollen Morphology of Embothrieae (Proteaceae)

This is the first of two papers detailing pollen morphology and evolution with the tribe Embothrieae comprising eight genera and ca. 56 spp. The present paper examines pollen of subtribes Buckinghaminae (Buckinghamia; 2 spp., Opislhiolepis; 1 sp.), Stenocarpinae (Strangea; 3 spp.), Stenocarpus (ca. 27 spp.) and Lomatiinae (Lomatia; ca. 12 spp.) in the light microscope and scanning and transmission electron microscopes. Pollen is medium-sized, oblate, foveolate to microreticulate to reticulate, and predominantly columellate with a complex modified postvestibulate aperture morphology. Pollen data indicate ties between Lomatia and Stenocarpus on the one hand and Stenocarpus and Strangea on the other. Though Buckinghamia and Opislhiolepis have been placed in the same subtribe, the unique combination of pollen features in each suggests only a remote relationship to each other as well as to remaining Embothrieae. Comparisons to the remaining genera of Embothriae (Embothrium, Oreocallis, Telopea) and overall analysis of pollen evolution within the tribe are detailed in the subsequent paper.     

Gaertnera and Pagamea: Genera Within the Psychotrieae or Constituting the Tribe Gaertnereae? A Wood Anatomical and Palynological Approach

The possible alliance between Gaertnera and Pagamea (Rubiaceae-Rubioideae), two genera from the Old and New World, respectively, is investigated on the basis of wood anatomy and pollen morphology. Nowadays, the main point of discussion about the taxonomic position of these genera is whether they belong to the Psychotrieae or constitute a tribe Gaertnereae characterised by their secondarily superior ovary and sheathing stipules. Both the wood and pollen of the genus pair are found to show specific features absent in other genera of the Psychotrieae, e.g. parenchyma bands in the xylem and endexine thickenings on the polar sites of the pollen apertures. Nevertheless Gaertnera and Pagamea share many other characters with the Psychotrieae. Wood and pollen convincingly demonstrate the very close affinity of the two genera. The sister pair differs in so many features from other Psychotrieae, that Gaertnera and Pagamea should constitute at least a subtribe Gaertnerinae, formally recognized here. The general lack of profound studies on the affinities within the very large tribe Psychotrieae makes further comments on the taxonomic relationships of the Gaertnerinae difficult.     

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.     

The production of floral oils byMonttea (Scrophulariaceae) and the function of tarsal pads inCentris bees

Plant species that secrete oil as their primary floral reward are rare and sporadically found in the angiosperms. We report here thatMonttea, a genus previously unsuspected of being an oil-plant, produces lipids from trichome elaiophores on the inside of the lower (anterior) lip. The discovery of the production of oils by species of this S. American genus explains the occurrence of unusual dual-function collecting structures in ArgentineCentris (Hymenoptera: Anthophoridae) and explains the presence of oil-collecting bees in regions where oil-secreting flowers were previously thought to be absent. The behavior of these centridine pollinators onMonttea flowers parallels that of oil-collecting bees onDiascia (Scrophulariaceae) in S. Africa.     

POLLEN MORPHOLOGY AND SYSTEMATIC RELATIONSHIPS AMONG NEW WORLD SPECIES IN TRIBE ANTIRRHINEAE (SCROPHULARIACEAE)

The morphology of mature pollen grains among 12 New World genera in tribe Antirrhineae has been examined with light and scanning electron microscopy. Pollen from 29 American species have mean polar diam that range from 17 to 26 μm, have a tectate structure with perforate, microreticulate, or reticulate surface pattern, and are subspheroidal to prolate, and trizonocolporate with fusiform or narrowly oblong colpi that are free or occasionally fused at the poles. Among the genera, pollen size is positively associated with style length but not haploid chromosome number. Three major morphological categories are recognized based on grain shape, exine structural pattern, and colpus shape. Two of these morphotypes each characterize individual genera (Mohavea, Linaria). The other morphological category includes ten New World genera and has been divided into three subcategories based on variation in lumina diam and intermittent occurrence of fused colpi. Only Antirrhinum and Pseudorontium have species or populations with pollen in more than one subcategory. Concordance of morphological, anatomical, chromosomal, and palynological data suggests that Linaria and Mohavea are best placed in separate subtribes from other native American genera. Compared to Old World taxa in tribe Antirrhineae, the most common pollen type (subspheroidal, microreticulate, apocolpate, fusiform colpi) may be the most primitive. Pollen grains with fused colpi (Antirrhinum p.p., Pseudorontium) and perforate (Mohavea) or reticulate (Holmgrenanthe) structural patterns probably represent derived types.     

Pollen-ovule patterns in tribeTrifolieae (Leguminosae)

Relative pollen and ovule production in the genera of the legume tribeTrifolieae is explored particularly as to howMedicago with its explosive pollination mechanism compares with its allies.Medicago produces much larger, although much fewer pollen per ovule than the other five genera in the tribe; this is interpreted as a consequence of its highly specialized, irreversible pollination mechanism, which allows only one effective exchange of pollen with pollinators.Melilotus andTrifolium produce a comparatively large quantity of pollen; this can be related to their floral characteristics requiring abundant pollen to achieve fertilization. InMedicago andTrigonella, annuals convert a higher proportion of pollen grains and ovules into seeds than perennials.     

Pollen morphology of the tribe Lithospermeae of Boraginoideae in China and its taxonomic significance

The pollen morphology in 15 species representing five genera in the tribe Lithospermeae of Boraginoideae (Boraginaceae) has been investigated and illustrated using light microscopy and scanning electron microscopy. The tribe Lithospermeae is very diverse in pollen morphology. The pollen grains are 10.4–41.8 × 7–33.1 μm in size with subspheroidal, prolate, cocoon, dumbbell, and ovoid shapes. The pollen apertures are of five types: 3-colporate, 3-syncolporate, 4-8-colporate, 4-6-syncolpate, and 6-7-colpate types. The exine ornamentations are generally smooth or rugulose, sparsely echinulate, and rarely rugulate. Based on pollen morphology, we developed a key to identify the genera of the tribe Lithospermeae, compared the pollen apertures among tribes of Boraginoideae, explained the evolutionary trends of the pollen grains, and discussed the taxonomic position of the tribe Lithospermeae. The palynological data suggest pollen of the eurypalynous type and support the proposal that the tribe Lithospermeae is in the primitive position of the subfamily Boraginoideae, and Echium Linn. is in the tribe Lithospermeae rather than in a novel tribe. Our observations have application potential for identification of pollen fossils of the tribe Lithospermeae.     

Morphology of pollen and orbicules in the tribe Spiraeeae (Rosaceae) and its systematic implications

This study provides pollen data for 38 representative taxa belonging to all nine genera in the current classification of the tribe Spiraeeae (Rosaceae) including the monotypic Korean endemic genus Pentactina, and considers the distribution of orbicules for the first time. Pollen morphology and wall stratification were investigated using light, scanning electron and transmission electron microscopy. Spiraeeae pollen grains are small to medium in size (P = 6.9–34.0 μm, E = 7.1–28.0 μm), oblate to prolate in shape (P/E = 0.66–1.48) and tri-colporate. Spiraeeae pollen is generally characterised by striate sexine ornamentation, but four ornamentation types are recognised based on the length and direction of the ridge patterns. The observed variation in sexine ornamentation is particularly valuable at the generic level. The exine stratification of the representative Spiraeeae studied is similar and characterised by unbranched columellae and a continuous endexine. Orbicules are present in three genera of the tribe (Luetkea, Sibiraea and Xerospiraea). Orbicule distribution patterns indicate that the absence of orbicules is a synapomorphic condition of the more derived clade, comprising Pentactina + Petrophytum + Kelseya + Spiraea.     

Structure,ultrastructure and evolution of floral nectaries in the twinflower tribe Linnaeeae and related taxa (Caprifoliaceae)

Linnaeeae is a small tribe of Caprifoliaceae consisting of six genera and c. 20 species. In Linnaeeae, floral nectaries are located on the corolla‐filament‐tube and nectar is produced from unicellular glandular hairs. We studied 23 taxa using scanning electron microscopy (SEM), light microscopy (LM) and transmission electron microscopy (TEM) and found two distinct nectary morphologies, zonate and gibbous types, and two distinct types of glandular hair, clavate and smooth base types. Plesiomorphic characters associated with the nectary and identified in the tribe include hypocrateriform corollas, dichogamous flowers, zonate nectaries, wet papillate stigmas, vestigial nectary disc and smooth pollen grains. Apomorphic characters include bilabiate corollas, homogamous flowers, bulging nectaries, dry papillate stigmas and echinulate pollen grains. The nectary structure is similar in Vesalea and Linnaea and differs from the rest of the tribe, in accordance with recent phylogenetic results. Nectar secretion is typically granulocrine with subcuticular accumulation of nectar, which we compared with the secretion in multicellular hairs of Adoxa moschatellina. The cuticle on the hair becomes detached from the cell wall and large subcuticular spaces filled with nectar are formed. Nectar is probably released in areas with a thin cuticle. In Zabelia, the smooth basal part of the hair could help to build up the hydrostatic pressure.     

On the pollen morphology of Scyphosyce and Dorstenia (Moraceae)

The pollen grains of the two species of Scyphosyce are alike and show distinct resemblances with those of several species of Dorstenia. Pollenmorphological data confirm the taxonomic suggestion to place Scyphosyce and Dorstenia together in the tribe Dorsteniae.     

SYSTEMATIC SIGNIFICANCE OF POLLEN NUCLEAR NUMBER IN EUPHORBIACEAE,TRIBE EUPHORBIEAE

Pollen nuclear number is determined in 139 species of 5 genera in the Euphorbieae, subtribe Euphorbiinae. The 111 new determinations are tabulated along with previous reports, and the results indicate that the distribution of binucleate (II) and trinucleate (III) pollen is strongly associated with the taxonomic groupings within the Euphorbieae. Although binucleate pollen is probably primitive within the tribe Euphorbieae, as suggested by the nuclear condition in Neoguillauminia, the situation in Euphorbia still requires further elucidation. Within Euphorbia, the morphologically most primitive species studied have III pollen despite the fact that II pollen is presumably the original condition for the subtribe Euphorbiinae. In Euphorbia, II pollen only is reported from nine sections and III pollen only from ten sections, while in four sections (Esula, Goniostema, Aphyllis, and Deuterocalli) both II and III pollen have been found. The New World species of Euphorbia nearly all have III pollen, whereas the vast majority of the African succulents have II pollen. The genera of New World origin, Chamaesyce and Pedilanthus, have III pollen, while the African genera Monadenium and Synadenium have II pollen. Independent derivations of III pollen from II pollen appear to have occurred in sections Goniostema, Aphyllis, and Deuterocalli (all of subg. Euphorbia). There is no evidence that reversals from III to II pollen have occurred.     

Pollen morphology and systematics of Atripliceae (Chenopodiaceae)

Pollen morphology of 58 species from 17 putative genera of the tribe Atripliceae (Chenopodiaceae) was investigated using light (LM) and scanning electron microscopy (SEM). Morphological variation was analyzed based on a dense sampling of the subtribes Atriplicinae and Eurotiinae, including many of the species in the two largest genera: Atriplex and Obione. The pantoporate pollen grains of Atripliceae are characterized by their spheroidal or subspheroidal shape, flat or moderately vaulted mesoporia with 21–120 pores, tectum with 1–8 spinules and 5–28(?38) puncta per?µm², and 1–13 ectexinous bodies bearing 1–7 spinules each. Taxonomic relevance of the most important pollen morphological characters is discussed (pollen diameter, pore number, pore diameter, interporal distance, spinule and puncta density and ratio, number of ectexinous bodies, and their spinules). Pollen morphological data support the exclusion of Suckleya from the tribe and the recognition of subtribe Eurotiinae, but suggest that it needs to be reviewed. Pollen does not support generic recognition of Atriplex, Neopreissia and Obione and infrageneric subdivisions as currently recognized, and suggests the need to review them. Smaller or monotypic genera, such as Axyris, Ceratocarpus, Endolepis, Krascheninnikovia, Microgynoecium, Proatriplex and Spinacia have distinctive pollen morphological characters that support their generic status. Grayia needs to be reevaluated; although its two species are distinct from all the other species in the study, there are notable differences between each of them, and this suggests they may not form a natural group. Multivariate techniques were employed to investigate if there are discrete patterns of variation within Atripliceae. Principal Component Analyses (PCA) weakly differentiates four groups based on variation in pore number, puncta density per?µm², and ratio between spinule and puncta density per?µm²; species of Ceratocarpus, Haloxanthium, Krascheninnikovia, Manochlamys, Microgynoecium, Spinacia, and some species of Atriplex and Obione are isolated. Preliminary results indicate that pollen data are potentially useful in the classification of the tribe, and further studies will be of taxonomic value.     

Pollen wall development in Malvastrum corchorifolium (Malvaceae)

The Malvaceae family has many species that are the subject of systematic controversy. Its broad spectrum of fruit and pollen morphology has challenged taxonomists because often morphological studies lead to different conclusions. Therefore, this study of selected stages of pollen wall development in Malvastrum corchorifolium is offered as another approach to verify its current taxonomic assignment and assist in the clarification of malvacean ancestry. Aperture number, endexine thickness and the presence of basal cushions are features that concur with its present placement in the Abutileae tribe, Sidinae subtribe and genus Malvastrum. Its advanced spine morphology, long spines with pointed apices and prominent basal cushions, suggest that Malvastrum corchorifolium is highly specialized and evolutionarily advanced.     

The systematic relationships ofAegialitis (Plumbaginaceae) as revealed by pollen morphology

Light and first SEM observations on the pollen ofAegialitis indicate that the two species of the genus are clearly distinguishable from all other genera of the tribeStaticeae to whichAegialitis presently is relegated. Intraspecific exine or aperture dimorphism which appears frequently in this tribe, is not recorded in this genus.A. annulata andA. rotundifolia have distinct monomorphic pollen and show a great resemblance in exine features with the tribePlumbagineae, particularly species ofPlumbago. Considering these and other features, separate subtribal status forAegialitis within thePlumbagineae is proposed.     

Veratrum album and V. nigrum (Melanthiaceae) in Italy: Micromorphology and systematics

ABSTRACT

Floral anatomy and pollen morphology of the two European species of Veratrum (V. nigrum and V. album subsp. lobelianum) from Italy are described in the context of the systematics of the genus and tribe Melanthieae (Melanthiaceae sensu stricto). Septal nectaries are absent, as in other Liliales. Most characters in Veratrum are typical of Melanthieae: e.g. semi-inferior ovaries, fused carpellary bundles and operculate pollen.     

Pollen and the Evolution of Arctotideae (Compositae)

Recent molecular studies have elucidated the phylogeny of Compositae tribe Arctotideae, and found it to contain two, well supported, monophyletic subtribes, Arctotidineae and Gorteriinae, as well as some polyphyletic and problematic genera. On the basis of this new information, it may now be possible to identify diagnostic characters and synapomorphies to support the groupings defined within Arctotideae. Pollen characters have been shown to be particularly variable in Compositae. This paper aims to investigate the utility of those characters in the context of recent molecular phylogenies, in order to determine synapomorphic and diagnostic characters in Arctotideae. The pollen of each genus is described, illustrated with scanning electron micrographs, and optimised on a phylogeny of the tribe. Many pollen characters were found to be very informative when considered in the context of the current best estimate of phylogenetic relationships. Pollen morphology provides synapomorphies for clades at a number of hierarchical levels within Arctotideae, including the two subtribes, Arctotidinae and Gorteriinae, the grouping of Eremothamnus and Hoplophyllum, and smaller clades. It also supports the exclusion of Platycarpha from the tribe. The plesiomorphic palynological state for the tribe is discussed. Particular attention is paid to the evolution of different patterns of lophae (surface ridges). A single origin for the lophate condition is proposed as the most parsimonious mode of evolution in Arctotideae.