Comparative floral structure and systematics in the balsaminoid clade including Balsaminaceae,Marcgraviaceae and Tetrameristaceae (Ericales)

Based on molecular phylogenetic studies, Balsaminaceae, Tetrameristaceae (including Pellicieraceae) and Marcgraviaceae form the strongly supported first branching clade in the asterid order Ericales. Marcgraviaceae and Tetrameristaceae were proposed to be closely related in pre‐molecular studies, but the systematic position of Balsaminaceae has been controversial for some time and a relationship with the other two families was never suggested in pre‐molecular/pre‐cladistic times. However, interfamilial relationships in the clade are still unclear because of conflicting phylogenetic hypotheses from molecular analyses. In order to assess the validity of these molecular hypotheses from a morphological point of view, the floral morphology, anatomy and histology of Balsaminaceae, Tetrameristaceae and Marcgraviaceae are comparatively studied in detail. In addition, earlier literature is reviewed. The monophyly of the balsaminoid clade is strongly supported by floral structure, and a series of potential floral synapomorphies is identified for the clade. Prominent features shared by the three families include broad and dorsiventrally flattened filaments, thread‐like structures lining the stomia of dehisced anthers, secretory inner morphological surfaces of the gynoecium, ovules intermediate between uni‐ and bitegmic, incompletely tenuinucellar ovules, fruits with persistent style and stigma, seeds lacking endosperm and several anatomical/histological traits. The families are also distinctive because the bracts and/or sepals are petaloid and nectariferous. Further, the floral structure supports a sister group relationship between Balsaminaceae and Tetrameristaceae rather than any of the other possible interfamilial relationships. These two families share a caducous calyx, post‐genital fusion/coherence of filaments and ovary surface, latrorse anther dehiscence, commissural carpel lobes and ovules with a thickened funiculus and a constricted chalazal region. The occurrence of these features in Ericales is discussed. Future structural studies in other ericalean lineages and additional molecular studies are needed to further test these features with respect to their systematic value for the balsaminoid clade. Some may turn out to be true synapomorphies, whereas others may be recognized as plesiomorphies, as they may be more widely spread in Ericales than currently thought. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 325–386.     

Conflicting phylogenies of balsaminoid families and the polytomy in Ericales: combining data in a Bayesian framework

The balsaminoid Ericales, namely Balsaminaceae, Marcgraviaceae, Tetrameristaceae, and Pellicieraceae have been confidently placed at the base of Ericales, but the relations among these families have been resolved differently in recent analyses. Sister to this basal group is a large polytomy comprising all other families of Ericales, which is associated with short internodes. Because there are more than 13 kb of sequences for a large sampling of representatives, a thorough examination of the available data with novel methods seemed in place. Because of its computational speed, Bayesian phylogenetics allows for the use of parameter-rich models that can accommodate differences in the evolutionary process between partitions in a simultaneous analysis. In addition, there are recently proposed Bayesian strategies of assessing incongruence between partitions. We have applied these methods to the current problems in Ericales phylogeny, taking into account reported pitfalls in Bayesian analysis such as model selection uncertainty. Based on our results we infer several, previously unresolved relationships in the order Ericales. In balsaminoid families, we find that the closest relatives of Balsaminaceae are Marcgraviaceae. In the Ericales polytomy, we find strong support for Pentaphylacaceae sensu APG II as the sister group of Maesaceae. In addition, Symplocaceae receive a position as sister to Theaceae and these families form a monophyletic group together with Styracaceae-Diapensiaceae. At the base of this clade are Actinidiaceae and Clethraceae. The positions of Ebenaceae and Lecythidaceae remain uncertain.     

Relationships within balsaminoid Ericales: a wood anatomical approach

Wood samples of 49 specimens representing 31 species and 11 genera of woody balsaminoids, i.e., Balsaminaceae, Marcgraviaceae, Pellicieraceae, and Tetrameristaceae, were investigated using light microscopy and scanning electron microscopy. The wood structure of Marcgraviaceae, Pellicieraceae, and Tetrameristaceae is characterized by radial vessel multiples with simple perforation plates, alternate vessel pitting, apotracheal and paratracheal parenchyma, septate libriform fibers, and the presence of raphides in ray cells. Tetrameristaceae and Pellicieraceae are found to be closely related based on the occurrence of unilaterally compound vessel-ray pitting and multiseriate rays with long uniseriate ends. The narrow rays in Pelliciera are characteristic of this genus, but a broader concept of Tetrameristaceae including Pelliciera is favored. Within Marcgraviaceae, wide rays (more than five-seriate) are typical of the genus Marcgravia. Furthermore, there is evidence that the impact of altitude and habit plays an important role in the wood structure of this family. The wood structure of Balsaminaceae cannot be compared systematically with other balsaminoids because of their secondary woodiness. Balsaminaceae wood strongly differs due to the presence of exclusively upright ray cells in Impatiens niamniamensis, the absence of rays in Impatiens arguta, and the occurrence of several additional paedomorphic features in both species.     

Palynological variation in balsaminoid ericales. I. Marcgraviaceae

BACKGROUND AND AIMS: Marcgraviaceae are a rather small family of seven genera and approx. 130 neotropical species. This study aims to present a detailed palynological survey of the family in order to comment on the intrafamily relationships and possible correlations with pollinators. METHODS: In total, 119 specimens representing 67 species and all genera are observed using light microscopy and scanning electron microscopy. Furthermore, eight species from five genera are studied with transmission electron microscopy. KEY RESULTS: Our results show that pollen grains of Marcgraviaceae are small (20-35 microm), have three equatorial apertures, granules on the colpus membrane, oblate spheroidal to prolate spheroidal shapes, mainly psilate to perforate ornamentations, and lalongate colpus-shaped thinnings at the inner layer of the exine, and show the presence of orbicules. Based on our fragmentary knowledge of the pollination biology of the family, there are no clear correlations between pollinators and pollen features. CONCLUSIONS: The genus Marcgravia has a high percentage of reticulate sexine patterns and a relatively thin nexine. Sarcopera can be defined by the presence of an oblate spheroidal to even suboblate shape, while Ruyschia and Souroubea typically show prolate spheroidal to subprolate pollen grains. The presence of a thick foot layer in the pollen wall is characteristic of the genera Norantea, Sarcopera and Schwartzia. Pollen features that are taxonomically useful within the family are the shape, sexine sculpturing, and ultrastructure of the pollen wall.     

Diversity and evolution of floral structure among early diverging lineages in the Ericales

This is a combination of review and original data on floral structure and diversity in the two earliest diverging lineages of the Ericales, i.e. the balsaminoids, comprising Balsaminaceae, Marcgraviaceae and Tetrameristaceae, and the polemonioids, comprising Fouquieriaceae and Polemoniaceae. Each clade is strongly supported in molecular studies, while structural synapomorphies have largely been lacking. For the balsaminoid families, we compare floral morphology, anatomy and histology among selected taxa and find that the entire clade is strongly supported by the shared presence of nectariferous tissue in the floral periphery, thread-like structures on anthers, truncate stigmas, secretion in the ovary, as well as mucilage cells, raphides and tannins in floral tissues. A possible sister group relationship between Balsaminaceae and Tetrameristaceae is supported by the shared presence of post-genital fusion of filaments and ovary and a star-shaped stylar canal. For polemonioids, we document unexpected diversity of floral features in Polemoniaceae, partly providing structural links to Fouquieriaceae. Features include cochlear and quincuncial corolla aestivation, connective protrusions, ventrifixed anthers and nectariferous tissue in the base of the ovary. In addition, we outline future directions for research on floral structure in the Ericales and briefly discuss the general importance of structural studies for our understanding of plant phylogeny and evolution.     

Pollen morphology in Campanulaceae IV

The exine structure has been studied in Campanulaceae s. lat. The results are combined with those in previous studies of shape, sculpturing and aperture conditions. Fifteen further species have been studied by SEM. Two main groups of pollen are found: (1) porate pollen with spinules and ridges/protrusions or a low relief reticulum, combined with an ektexine varying from simple to complex and a lamellated endexine; (2) 3–colpate/colporate pollen with a high relief reticulate/striate surface sculpturing, in general a homogeneous ektexine and an endexine lacking lamellae. The first group corresponds to Campanulaceae s. str., the second to Lobeliaceae s. str. Some genera like Cyananthus, Codonopsis and Parishella have unique characters which make them difficult to place in any of the two main groups.     

Pollen morphological support for the Catesbaeeae-Chiococceae-Exostema-complex (Rubiaceae)

The pollen morphology of the Catesbaeeae-Chiococceae-Exostema complex as recently treated by Delprete (1966) was examined with LM and SEM. The group is remarkably stenopalynous; typical representatives have medium sized, 3-colpate pollen with a perforate tectum covered with microspines. The inner nexine ornamentation is pronounced and offers more variation than the sexine pattern. A typology of the inside structures is presented based on LM observations and SEM observations of sectioned grains. Orbicules are common and numerous in the Catesbaeeae and Exostema-group; for most genera of the Chiococceae confirmation is needed of orbicule presence. All orbicules observed are relatively large (1-4 μm) and spiny. Pollen and orbicule morphology proved to be a powerful tool to delimit the Catesbaeeae-Chiococceae-Exostema complex. The overall delimitation of the complex is corroborated with our pollen data. The genera Mastixiodendron and Placocarpa, however, can be excluded from the complex based on their pollen morphology. Mastixiodendron has 3-colporate, perforate pollen without microspines and the endocolpi are fused into an endocingulum. Pollen of Placocarpa is reticulate and 3-colporate with perpendicular endocolpi.     

The multiple fuzzy origins of woodiness within Balsaminaceae using an integrated approach. Where do we draw the line?

Background and Aims

The family Balsaminaceae is essentially herbaceous, except for some woodier species that can be described as ‘woody’ herbs or small shrubs. The family is nested within the so-called balsaminoid clade of Ericales, including the exclusively woody families Tetrameristaceae and Marcgraviaceae, which is sister to the remaining families of the predominantly woody order. A molecular phylogeny of Balsaminaceae is compared with wood anatomical observations to find out whether the woodier species are derived from herbaceous taxa (i.e. secondary woodiness), or whether woodiness in the family represents the ancestral state for the order (i.e. primary woodiness).

Methods

Wood anatomical observations of 68 Impatiens species and Hydrocera triflora, of which 47 are included in a multigene phylogeny, are carried out using light and scanning electron microscopy and compared with the molecular phylogenetic insights.

Key Results

There is much continuous variation in wood development between the Impatiens species studied, making the distinction between herbaceousness and woodiness difficult. However, the most woody species, unambiguously considered as truly woody shrubs, all display paedomorphic wood features pointing to secondary woodiness. This hypothesis is further supported by the molecular phylogeny, demonstrating that these most woody species are derived from herbaceous (or less woody) species in at least five independent clades. Wood formation in H. triflora is mostly confined to the ribs of the stems and shows paedomorphic wood features as well, suggesting that the common ancestor of Balsaminaceae was probably herbaceous.

Conclusions

The terms ‘herbaceousness’ and ‘woodiness’ are notoriously difficult to use in Balsaminaceae. However, anatomical observations and molecular sequence data show that the woodier species are derived from less woody or clearly herbaceous species, demonstrating that secondary woodiness has evolved in parallel.     

Pollen morphology of endemic species of the Horton Plains National Park,Sri Lanka

The pollen morphology of 27 endemic and palaeoecologically-important species belonging to 16 families and 20 genera from the mountain rain forests of Sri Lanka was studied using both light (LM) and scanning electron microscopy (SEM). The pollen grains exhibit a wide range of morphological characters. Their sizes (P and E) are in the range 10-53 2 9-60 mum, and they are distributed among seven shape classes. All the taxa have radially-symmetrical pollen grains except for Impatiens (bilateral). The amb varies from rounded to triangular, with intermediate shapes possible. The pollen grains of all the Lauraceae species studied are inaperturate and microspinose, with a thin, fragile, perforated exine. Sarcococca zeylanica (Buxaceae) has pantoporate pollen grains and a Croton -pattern consisting of faintly striate pegs of various shapes attached or enveloping the smooth rings. Colp(or)ate, syncolpate to parasyncolp(or)ate pollen grains with rugulate to perforate exine patterns occur in Eugenia mabaeoides ssp. mabaeoides and Syzygium ssp. (Myrtaceae). Osbeckia walkeri (Melastomataceae) has heterocolpate pollen with fossulate, foveolate to perforate exine pattern. 3-colporate, microreticulate pollen grains are met with in Euonymus revolutus (Celastraceae), and scabrate-microreticulate ones in Calophyllum walkeri (Clusiaceae). Hedyotis lawsoniae (Rubiaceae) pollen is microreticulate to perforate, while pollen of the Elaeocarpus (Elaeocarpaceae) species is 3-colporate, smooth to scabrate-perforate. Casearia thwaitesii (Flacourtiaceae) has 3-colporoidate pollen grains with a smooth, imperforate to occasionally sparsely perforate exine. Mastixia spp (Cornaceae) have a granular-perforate exine pattern, while Adinandra lasiopetala (Theaceae) has a smooth-perforate exine pattern. Triangular 3 (4)-colporate, and indistinctly faintly microreticulate-perforate pollen was found in Rhamnus arnottianus (Rhamnaceae), rugulate-perforate in Isonandra montana (Sapotaceae) and verrucate-perforate in Symplocos elegans (Symplocaceae). Glochidion coriaceum (Euphorbiaceae) has a 5-colporate grain with a coarsely reticulate exine pattern. Pollen grains of Impatiens walkeri and I. thwaitesii (Balsaminaceae) are bilateral, 4-colpate, reticulate and granular. A pollen key based on light microscope is constructed for all the taxa studied.     

中国紫葳科花粉形态的研究

对紫葳科18属27种花粉进行了光镜和扫描电镜的观察,其中有些种是第一次观察或报道。紫葳科花粉明显地属多型花粉,萌发孔类型和纹饰特征多种多样。萌发孔基本上可分为3（－4）沟,3（－4）孔沟,3孔,多沟及散沟等类型;纹饰则可分为皱波状饰,穴状纹饰,穴－网状纹饰,网状纹饰,刺状纹饰以及表面光滑等。值得注意的是在菜豆树属Rademoachera Zoll.et Moritzi中,菜豆树R．sinica和滇菜豆树R.yunnanensis虽为同一个属,但其萌发孔类型和纹饰特征截然不同,菜豆树花粉为3孔,具网状纹饰;而滇菜豆树花粉为3孔沟,具穴状纹饰。就花粉形态学而言,紫葳科也许是多元发生的。     

Phylogenetic relationships in the order Ericales s.l.: analyses of molecular data from five genes from the plastid and mitochondrial genomes

Phylogenetic interrelationships in the enlarged order Ericales were investigated by jackknife analysis of a combination of DNA sequences from the plastid genes rbcL, ndhF, atpB, and the mitochondrial genes atp1 and matR. Several well-supported groups were identified, but neither a combination of all gene sequences nor any one alone fully resolved the relationships between all major clades in Ericales. All investigated families except Theaceae were found to be monophyletic. Four families, Marcgraviaceae, Balsaminaceae, Pellicieraceae, and Tetrameristaceae form a monophyletic group that is the sister of the remaining families. On the next higher level, Fouquieriaceae and Polemoniaceae form a clade that is sister to the majority of families that form a group with eight supported clades between which the interrelationships are unresolved: Theaceae-Ternstroemioideae with Ficalhoa, Sladenia, and Pentaphylacaceae; Theaceae-Theoideae; Ebenaceae and Lissocarpaceae; Symplocaceae; Maesaceae, Theophrastaceae, Primulaceae, and Myrsinaceae; Styracaceae and Diapensiaceae; Lecythidaceae and Sapotaceae; Actinidiaceae, Roridulaceae, Sarraceniaceae, Clethraceae, Cyrillaceae, and Ericaceae.     

国产爵床科芦莉花族植物的花粉形态

报道了国产爵床科Acanthaceae芦莉花族Ruellieae芦莉花亚族Ruelliinae 2属7种、假杜鹃亚族Barlerinae 1属3种和马蓝亚族Strobilanthinae 16属34种植物扫描电镜下的花粉形态.芦莉花亚族的地皮消属Pararuellia和喜花草属Eranthemum的花粉均为圆球形,具3孔或3孔沟,外壁为不同的网状结构; 假杜鹃亚族的假杜鹃属Barleria的花粉为长球形,具3孔沟,外壁亦为网状结构;马蓝亚族植物(包含广义的马蓝属Strobilanthes s.l.)花粉形态多样,结构复杂.依据花粉萌发孔和外壁纹饰特征,可将马蓝亚族16属植物和上述两亚族3属植物的花粉形态归纳成3大类型: 1. 具3孔类型.其中又有(1)外壁具网状纹饰者,见于地皮消属; (2)外壁具芽胞状纹饰者,见于黄猄草属Championella; (3)外壁具刺状(棒状)纹饰者,见于南一笼鸡属Paragutzlaffia、叉花草属Diflugossa和假蓝属Pteroptychia.2. 具3孔沟及具3孔沟与假沟类型(肋条带型).其中又有(1)具3孔沟和网状纹饰者,见于喜花草属和假杜鹃属; (2)具刺状(棒状)纹饰者,见于南一笼鸡属、叉花草属和假蓝属; (3)具3孔沟与假沟,外壁纹饰具节隔、肋条带状或网状,网眼纵向排列成行,网眼内有细网纹者,见于耳叶马蓝属Perilepta、马蓝属Pteracanthus(大部分)、金足草属Goldfussia、紫云菜属Strobilanthes(部分)和合页草属Sympagis; (4)具3孔沟与假沟类型,肋条带状,但不具节隔,外壁纹饰网状,网眼不成行或不明显纵向排列,网内无细网纹者,见于尖蕊花属Aechmanthera、板蓝属Baphicacanthus、马蓝属(部分)和糯米香属Semnostachya; (5)具双脊及细网状纹饰者,见于环毛紫云菜Strobilanthes cycla.3. 具(4-)5孔沟及假沟类型(肋条带型),外壁具网状或拟网状纹饰,见于腺背蓝属Adenacanthus.另外兰嵌马蓝属Parachampionella、山一笼鸡属Gutzlaffia和肖笼鸡属Tarphochlamys的花粉有无萌发孔尚不清楚,有待进一步研究.综上所述,芦莉花族植物的花粉形态具有较高的多样性,是重要的分类性状.利用花粉形态特征能较好地区分高级分类群如亚科、族以及亚族,有时也有助于阐明类群之间的相互关系,甚至也能用于区分属、种和阐明其关系.     

Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological Significance

Pollen morphology of selected taxa from the Hengduan Mountains, Southwest China are investigated in the present study. Forty eight taxa (flowering in the summer, 46 species and 2 varieties) of 43 genera belonging to 27 families were observed by light and scanning electron microscopy. Pollen morphology is described in detail. Two pollen shapes are mainly found in these species: subprolate to prolate (271%) and prolate (25%). Spheroidal, subspheroidal to subprolate, suboblate and tetrad shapes can also be found in some species. Aperture type is mostly tricolporate, with a percentage of 438, and also contains tricolpate, pantoporate, stephanocolpate, 3 colporoid, 6 colpate, monocolpate. The most common ornamentation is finely reticulate, with a percentage of 50. Other exine ornamentations, such as coarsely reticulate, finely granulate, smooth, spinulose are also observed. The palynological documentation of these species will provide the modern palynological basis for paleopalynological studies of the Quaternary Strata. On the basis of pollen morphology characters, identification features of some species in the same families were discussed according to the results of cluster analysis used by SPSS. The ecological significance was also discussed based on the ecological properties of these taxa.     

The remarkable genus<Emphasis Type="Italic"> Coptosapelta</Emphasis> (Rubiaceae): pollen and orbicule morphology and systematic implications

The pollen morphology and distribution of orbicules were investigated in ten species of the genus Coptosapelta (Rubiaceae) using light, scanning electron and transmission electron microscopy. In general, Coptosapelta pollen is three- to five-pororate and suboblate to oblate-spheroidal. The sexine ornamentation varies from psilate to reticulate, and the pollen-wall stratification lacks columellae. Droplets are observed on the inner nexine surface. Distinct orbicules are found in five species. Orbicules are often lobed and appear to be aggregated and embedded in the tapetal remnants of the locule wall. Ultrastructurally, a single, flattened core is frequently observed. Coptosapelta has a unique pollen type within Rubiaceae and does not resemble possibly related genera. Pollen characteristics provide additional evidence supporting earlier findings that Coptosapelta represents an isolated lineage in Rubiaceae.     

Pollen morphology of Adenandra (Rutaceae: Diosminae) and its taxonomic implications

The pollen morphology of Adenandra Willd. (Rutaceae: Diosminae) was investigated to determine its taxonomic significance. Pollen of 27 of the 30 infrageneric taxa (representing 16 of the 18 species) was investigated by LM, SEM and TEM. Adenandra differs from all other Diosminae in having 4-colporate rather than 3-colporate pollen grains. This supports the alleged monophyletic status of the genus. Exine morphology and structure, however, is extremely diverse considering the size of the genus, with eight distinct pollen types and four subtypes being discerned. In some members the grains have uniform macroreticulate, striate or striato-reticulate sculpturing, whereas in others the poles have striate, striato-reticulate or reticulate-perforate sculpturing with various types of reticulate or rugulate sculpturing at the mesocolpia. Taxonomic groupings revealed by the pollen characters suggest elationships between taxa that were not previously apparent. It is suggested that evolutionary diversification among species of Adenandra is often more strikingly reflected by pollen morphology than by macromorphology, hence the eurypalynous state of the genus.     

Pollen morphology of the subtribe Cuspariinae (Rutaceae)

The neotropical subtribe Cuspariinae (Rutaceae) comprises as many as 26 genera and over 125 species. Pollen grains from 111 collections representing 71 species and 24 genera were examined by LM, SEM, and TEM. The pollen morphology of this subtribe is very diverse. Grains are mostly 3–6-aperturate and colporate, rarely porate (Spiranthera) or pantocolporate (Almeidea). Exine sculpturing is most commonly reticulate, sometimes perforate, foveolate-perforate, foveolate, foveolate-reticulate, reticulate, striate-reticulate, echinate, clavate, or baculate. The exine structure is columellate and tectate-perforate, columellate and semitectate, or intectate and is stratified into ektexine and endexine. The exine ofLeptothyrsa is distinctive in that the ektexine of the mesocolpium is longitudinally deeply ridged. The pollen ofHortia, characterized by a psilate exine with rare perforations, a very thick foot-layer, and reduced columellae, is unlike that of any member of the Cuspariinae and offers no support for the transfer of this genus from the Toddalioideae. The pollen data correlate with macromorphological characters and are taxonomically useful.     

Palynological and chemical volatile components of typically autumnal honeys of the western Mediterranean

Measurements and observations of pollen grains of varying structure before and after dehiscence of the anther sac illustrate large magnitudes of harmomegathic changes in volume and shape. Such changes are compared for colpate, porate, and colporate grains, and it is suggested that the nature of the harmomegathal action may serve to distinguish colpi or furrows from pores. Consideration of the requirement to allow harmomegathy while preventing wall collapse may provide at least partial explanation for evolution of the internal structure and external sculpturing of pollen grain walls.     

Study on the Pollen Morphology of the Chinese Family Oleaceae

This paper deals with the pollen grains of 11 genera, 22 species in Chinese Oleaceae. A comparative study on the pollen grains was carried out by means of scanning electron microscope and light microscope. The pollen grains of the family Oleaceae are almost spheroidal, oblate and prolate, in polar view tlie majority of the pollen grains are tri-lobulate-circular, the minority are tetra-lobulate-circular and dilobulate-circular. Pollen size: (15-65) × (12- 60)μ, 3-colporate and 3-colpate the minority of the pollen grains 4-colpate and 2- colpate colpus usually distinct and the en doaperature vague. The exine oonsists of two layers, the sexine thicker than the nexine, with big reticulum or fine reticulum, the majority of the reticulum are very distinct. On the basis of the pollen size and exine ornamentation, the pollen grains are divided into following two main types: (1) The larger pellen grains and the exine with big reticulum are found in the genera: Jasminum, Nyctanthes, Ligustrum, and Syringa. (2) The smaller pollen grains and the exine with finer reticulum are found in the genera: Osmanthus, Olea, Chionanthus, Fraxinus, Forsythia, Fontanesia and Linociera. In view of the features of pollen morphology, this family is much similar to the family Itamamelidaeeae. For example, the first pollen type resembles the genus Coryiopsis of the family Hamamelidaceae, but there are some differences between them, the former with a bigger reticulate exine and without any colpal membrane, the latter with a finer reticulate apocolpium and eolpate membrane. The second pollen type is much similar to that of the genera Hamamelis and Loropetalum of family Hamamelidaceae.     

POLLEN STUDIES IN THE GENUS CORDIA (BORAGINACEAE)

The pollen of 40 species of Cordia L. was studied by light microscopy and scanning and transmission electron microscopy. Three major types were found, 3-porate with a reticulate sexine, 3-colpate to 3-colporoidate with a striatoreticulate sexine, and 3-colpate to 3-colporate with a spinulose sexine. These pollen types are correlated with significant differences in the inflorescence, flower, and fruit structures found in the sporophyte. The following revisions are suggested and discussed: that Cordia sect. Varronia P. Br. be elevated to generic status; that Cordia sebestena L. and a few close allies be treated as generically distinct; and that Cordia myxa L. be conserved as the lectotype for the genus, thus continuing a wide application of the epithet Cordia.