POLLEN APERTURE EVOLUTION AMONG THE SUBFAMILIES PERSOONIOIDEAE,SPHALMIOIDEAE, AND CARNARVONIOIDEAE (PROTEACEAE)

Pollen apertures were analyzed among the subfamilies Persoonioideae (seven genera; ca. 95 spp.), Sphalmioideae (one genus; one spp.), and Camarvonioideae (one genus; two spp.). Pollen was examined by light microscopy, cryosection, and transmission electron microscopy. Completed studies of pollen apertures among Grevilleoideae (ca. 40 genera; ca. 800 spp.), one of two major subfamilies in Proteaceae, provide a basis for comparison and analysis of aperture evolution among these subfamilies. Aperture characters within Persoonioideae are unique among Proteaceae examined to date. Five distinct aperture types occur among the three subfamilies, three of which (Placospermum, Persoonia, Bellendena) are restricted to Persoonioideae. Sphalmioideae and Camarvonioideae each exhibit a unique aperture organization. The most primitive aperture organization, and one unique to Placospermum, exhibits three main features: 1) a thin, granular endexine continuous around the grain; 2) a heterogeneous foot layer throughout the grain with increased disruptions at the aperture; and 3) only slight differences in exine characters between apertural and nonapertural regions. The Persoonia aperture type represents the next stage of aperture evolution which involves loss of endexine, restriction of a heterogeneous foot layer to the aperture, and marked differences in exine characters between apertural and nonapertural regions. The uniformly homogeneous ektexine in both nonapertural and aperture regions in Bellendena has developed independently. Sphalmium exhibits a primitively thin granular endexine though the restriction of endexine to the aperture is a derived condition. Carnarvonia exhibits several pollen characters also found among Grevilleoideae including: 1) a homogeneous nonapertural ektexine; 2) a slightly heterogeneous apertural ektexine; 3) a lamellate/granulate endexine organized into irregularly shaped “clumps” clustered around the aperture; and 4) a clear demarcation between apertural and nonapertural exine. These characters support the hypothesis that Carnarvonia may have diverged early from the pre-Grevilleoids.     

Pollen Morphology of the Family Magnoliaceae

The pollen morphology of the following genera is described: Manglietia (1 sp.), Magnolia (10 spp.), Talauma (2 spp.), Alcimandra (1 sp.), Michelia (5 spp.) and Liriodendron (2 spp.). It is shown that from a pollen morphological point of view the Magnoliaceae are a homogeneous, natural group.     

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 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 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.     

POLLEN MORPHOLOGY IN THE GENERA MONTTEA AND MELOSPERMA (SCROPHULARIACEAE)

Pollen of the scrophulariaceous genera Melosperma and Monttea was studied with light and scanning electron microscopy. Both have single, radiosymmetric, isopolar, trizoniaperturate pollen grains with usually ruptured colpus membranes, tectate-perforate to semitectate exines with a columellate interstitium and pitted or perforate nexines. Intergeneric differences in pollen morphology are established based on the differential correlation of sculpturing characters. Comparisons of the pollen of Melosperma and Monttea with that of the tribe Minuleae sensu stricto reveals no difference in the case of Melosperma and the existence of intermediate morphologies and “reticulate” similarities in the case of Monttea; the taxonomic significance of this morphological intergradation is considered. The transfer of Oxycladus aphyllus to Monttea is consistent with the pollen evidence, but proposed affinities between Monttea and tribe Antonieae of the Loganiaceae are not supported.     

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 MORPHOLOGY AND SYSTEMATICS OF SIPHONOGLOSSA SENSU LATO (ACANTHACEAE)

Pollen morphology of 15 species of Siphonoglossa and of two closely related groups was investigated. Two tribal-specific pollen types are found within Siphonoglossa sensu lato suggesting that the genus is artificial, composed of taxa belonging in several genera among two tribes (subtribes sensu Bremekamp) of Acanthaceae. Five taxa currently included in an informal subgeneric category of Siphonoglossa have tricolporate, prolate pollen (termed Type I) that is characteristic of Odontonemeae (= Odontoneminae, Justicieae). Pollen of the remaining taxa, belonging in two formal sections of the genus, are mostly 2-porate, bilateral (Type II) with a sexine sculpturing characteristic of Justicieae (= Justiciinae). Pollen of section Siphonoglossa is rather uniform, 2-porate, bilateral with lolongate pores, and seem to delimit a natural group. Taxa of section Pentaloba have a more heterogeneous pollen morphology, mostly 2-porate, bilateral with lalongate pores. Controversial aspects of the interpretation of pollen morphology in Justiciinae are presented and their relevance to this study are examined. Hypothetical trends in the evolution of pollen of Justiciinae are discussed and the application of pollen morphology to taxonomy of the genus is presented, including a recommendation for narrowing the generic concept of Siphonoglossa to the taxa of the type section.     

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.     

须芒草族植物花粉形态的观察

在光学显微镜下和扫描电镜下对禾本科须芒草族（Ａｎｄｒｏｐｏｇｏｎｅａｅ）中分隶于８个亚族３４个属的３６种植物的花粉进行形态面容和比较研究。结果显示,本族植物花粉形态较为一致,花粉近形或扁球形,单萌发孔,孔,周围加厚,具盖,外壁表面散布有颗粒。这表明其是一个自然类群。总体来讲,芬烨大的演化分异,只是表面纹饰的和芬烨大小有一;定的差异。纹饰可分为三种类型粗糙型,不明显疠状突起才明显疠状突起型。花粉开矿     

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 MORPHOLOGY OF THE CHLORAEINAE (ORCHIDACEAE: DIURIDEAE) AND RELATED SUBTRIBES

Pollen organization and morphology of the South American Chloraeinae (Orchidaceae) was examined by scanning electron microscopy and compared with that of the remainder of the otherwise Australasian Diurideae. All five genera of the Chloraeinae, Bipinnula, Chloraea, Codonorchis, Gavilea, and Geoblasta, and at least one genus from each of the other subtribes were sampled. The Australasian Diurideae are diverse in pollen organization and morphology. The two genera of the Acianthinae, Corybas and Acianthus, have very different pollen and their classification is questioned. Monad pollen organization of Pterostylis (Pterostylidinae) is reinterpreted as primitive and not secondarily derived. Pollen of the Chloraeinae is uniform in exine morphology and organization. Most species sampled have reticulate pollen which tends to be foveolate distally. The basic pollen unit of all Chloraeinae is the tetrad, except Codonorchis which possesses monads. Pollen morphology and organization of the Chloraeinae is most similar to the Caladeniinae, which supports the contention that the Chloraeinae including Codonorchis should be retained in the Diurideae.     

Pollen Morphology and Epidermal Characters of Leaves in Convallarieae (s. l.)

The tribe Convallarieae (sensu Krause 1930) consists of 7 genera, i.e. Convallaria, Speirantha, Reineckia, Theropogon, Tupistra, Rohdea and Aspidistra, but now generally recognized as two tribes, Convallarieae (the former 4 genera) and Aspidistreae (the rest). Observed in this work were pollen morphology of 17 species and epidermal characters of leaves of 12 species. All the 7 genera are covered in observations. Pollen grains in Convallarieae (s. str.) are all monosulcate and boat-shaped (Plate 1: A-F). The exine is rather uniformly microperforate (Plate 1: A-F); only Theropogon is exceptional in this respect: it has rugulate exine (Plate 1: O, P). Tang and Zhang (1985) have pointed out the heterogeneity of Theropogon in this tribe. Pollen morphology in the tribe Aspidistreae is widely variable. The genera Tupistra and Rohdea were shown to have monosulcate and boat-shaped pollen grains. Their exine is perforate or reticulate (Plate 1: G-N). Pollen grains in the genus Aspidistra, however, are nonaperturate and spheroidal. The exine in the genus varies from crass-rugulate, variously gemmate to tuberculate-baculate (Plate 2; A-H). The pollen morphology of Aspidistra is therefore distinctly different from that of Tupistra and Rohdea, which supports the Nakai's (1936) establishment of the tribe Rohdeae for Tupistra and Rohdea. Therefore, Krause's Convallarieae is reasonably divided into at least three tribes, Convallarieae (Speirantha, Convallaria, Reineckia and Theropogon), Aspidistreae (Aspidistra) and Rohdeae (Rohdea and Tupistra). The pollen characters of all the 7 genera are shown in Table 1. The evolutionary trends of pollen morphology (aperture and exine) in the three tribes are discussed and our major view-points are shown in Fig. 1. Observations on epidermal characters of leaves show that in the Convallarieae (s. 1.) stomatal apparatuses are all anomocytic; cuticular layer on the upper epidermis is mainly striatethickened or rather uniformly thickened (Plate 2: J--P; Plate 3: A-C, F-N), whereas in the genus Convallaria the cuticular layer is squamosely thickened (Plate 2: I; Plate 3: D, E).The epidermal characters of leaves in the 7 genera are summarized in Table 2.     

Pollen morphology and evolution in the genus Orobanche L. s.l. and its allied genera (Orobancheae/Orobanchaceae) in Turkey

The pollen morphology of 120 samples of the tribe Orobancheae, representing four genera and 40 species of Cistanche, Diphelypaea, Orobanche and Phelipanche native to Turkey, has been studied by light and scanning electron microscopy. Pollen of the tribe Orobancheae is typically isopolar, radially asymmetrical, oblate spheroidal or prolate and belongs to one of the following basic pollen types: inaperturate, tricolpate and syncolpate. Also, pollen heteromorphism is widespread among pest species of Orobanche and Phelipanche. Pollen characters display considerable variation among genera and species, but some characters are significant at the levels of genera and sections. The results of the SEM study show that there are various exine ornamentation types in these genera. The obtained results support the division of the traditionally circumscribed genus Orobanche sensu lato into two genera, Phelipanche and Orobanche. The pollen morphological features are discussed on the basis of molecular phylogeny of the taxa.     

Pollen morphology in Cytisus (Papilionoideae,Leguminosae) from Morocco and the Iberian Peninsula

Although it is generally accepted that the tribe Genisteae has rather homogeneous pollen, we have found considerable variation in pollen size and morphology in Cytisus and related genera (Argyrocytisus and Chamaecytisus), which match taxonomic groups defined on morphological grounds. The results of Principal Component Analysis of pollen characteristics show the following: Cytisus Sect. Alburnoides is well-delimited, with the smallest pollen grains and the simplest pattern in the tectum, fossulate-perforate to perforate; No separation of C. villosus (Sect. Cytisus) from Sect. Alburnoides is observed; Cytisus Sect. Spartopsis, with the largest pollen grains and reticulate to reticulate-fossulate ornamentation, forms an isolated group; C. fontanesii (Sect. Heterocytisus) stands apart from other Cytisus species by its homogeneous ornamentation and the perforate instead of psilate-punctate margo. Thus, the pollen data supports its transfer to the separated genus Chronanthos; Pollen ornamentation and morphology also support the separation of Argyrocytisus battandieri and Chamaecytisus mollis from Cytisus.     

Taxonomic significance of pollen types in the Guyana Highland-centred composite genera of Mutisioideae (Asteraceae)

The Guyana Highland-centred genera, or Stenopadus group, are a complex of species that belong to the tribes Mutisieae and Stifftieae of Asteraceae. The pollen morphologies of 29 species, from 12 of 13 genera of this complex, are described and illustrated using light and scanning electron microscopy. The exine sculpture and structure are highly significant. Four exine types, previously characterized, were distinguished: Gongylolepis , Wunderlichia , Mutisia , and Stenopadus . The characterization of the Stenopadus exine type is enlarged here. These exine types led to the recognition of four well-defined pollen types, whereas the spine length and exine thickness characterized six subtypes. Pollen types circumscribe genera or groups of genera, and some subtypes distinguish species. The pollen morphology within the complex is discussed in relation to the rest of Mutisioideae and other palynologically allied tribes of Cichorioideae. There is little correlation between pollen types and tribes; only the Stenopadus exine type is exclusive to the Stifftieae tribe. The remaining types are shared by the two tribes of the complex. Pollen morphology supports the hypothesis that this group of genera is close to the Gochnatia complex and the Cardueae tribe.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 327–340.     

POLLEN MORPHOLOGY AND DETAILED STRUCTURE OF FAMILY COMPOSITAE,TRIBE CICHORIEAE. I. SUBTRIBE STEPHANOMERIINAE

A survey of pollen morphology in 20 species representing the 11 genera of the North American subtribe Stephanomeriinae by light, scanning electron, and transmission electron microscopy revealed 10 of the 11 genera to have echinate, tricolporate pollen grains, Lygodesmia being the only genus with echinolophate pollen. Sectioned exines of most of the species examined are similar, being composed of ektexine and endexine. The ektexine surface is composed of spines which typically have globose perforate bases. A cavus occurs as a separation between the basis (foot layer) and the columellae in all of the genera examined except Chaetadelpha. Pollen of the two species of Glyptopleura were found to be strikingly different in exomorphology. Pollen of the putatively self-fertile G. marginata has much shorter spines than the closely related G. setulosa. Atrichoseris, Anisocoma, Calycoseris, Glyptopleura, Pinaropappus, Prenanthella, and most species of Malacothrix have pollen which lack paraporal ridges. The remaining genera, Chaetadelpha, Lygodesmia, Rafinesquia, and Stephanomeria have well-developed ridges of fused spine bases around the apertures. Pollen characters, particularly those of the aperture region, have been found to be systematically useful in the subtribe, therefore acetolyzed material gives more useful information than untreated pollen.     

POLLEN MORPHOLOGY OF THE EUCHARIDEAE (AMARYLLIDACEAE)

Eucharis, Caliphruria, and Urceolina form a monophyletic group of petiolate-leaved, Neotropical Amaryllidaceae ecologically specialized to the understory of primary tropical rain forest below 2,000 m elevation. Pollen morphology of the three genera is surveyed. Pollen grains of all species of Eucharis, Caliphruria, and Urceolina are boat-shaped elliptic, monosulcate, heteropolar, and bilateral in symmetry. Exine sculpturing is semitectate-columellate and reticulate in all species examined. A transformation series in reticulum coarseness and pollen grain size is described. The large pollen grain with coarse reticulum of most Eucharis species is considered ancestral. The fine reticulation of Caliphruria is considered derived and the exine morphology of Urceolina is intermediate. Both of these genera have medium-sized pollen grains. Exine dimorphism common to all Urceolina, but rare in Eucharis and Caliphruria, may be symplesiomorphous among those taxa exhibiting this morphology. The three genera are largely uniform in pollen grain ultrastructure, with completely ektexinous exines. Pollen grain size in Eucharis is not closely correlated with style length. Several wide-ranging species show considerable intraspecific variation in pollen size. Parallelisms in pollen grain evolution among related tribes of Neotropical Amaryllidaceae are discussed.     

POLLEN MORPHOLOGY OF THE LENNOACEAE

Drugg , Warren S. (California Res. Corp., La Habra.) Pollen morphology of the Lennoaceae. Amer. Jour. Bot. 49(10): 1027–1032. Illus. 1962.—The family Lennoaceae contains 3 genera: Pholisma, Lennoa, and Ammobroma. The pollen of Pholisma is 4-colporate with less common 3- and 5-colporate grains. Lennoa pollen is 3-colporate with rare 4-colporate examples. All Ammobroma pollen is 3-colporate. Pollen of both Lennoa and Ammobroma exhibits sexine thinning in the mesocolpia with resultant depressions. This feature is lacking in Pholisma. Sculpture is reticulate-simplibaculate on pollen of Lennoa and Pholisma, and reticulate-duplibaculate on pollen of Ammobroma. Pollen morphology supports placement of Lennoaceae in the Polemoniales near the Boraginaceae and Hydrophyllaceae.     

Pollen morphology of the tribe Cynoglosseae of Boraginoideae (Boraginaceae) in China

Pollen morphology of 16 Chinese species representing 7 genera in the tribe Cynoglosseae of Boraginoideae(Boraginaceae) was examined under LM and SEM, and 5 species under TEM. Pollen grains are cocoon-shaped, rarely subprolate, prolate or ovoid, very small, 7～ 15.7μm× 3.5 ~ 13.9 μm in size, P/E = 1.6～2.02; 3-colporate apertures alternate with 3-pseudocolpi, with equatorial endocingulus except those in Bothriospermum; exine surface is usually smooth, with or without perforations in two poles, rarely with tuberculate ornamentation; exine is rather thin and includes ectexine and endexine, while ectexine consists of imperforate tectum, columellae and foot-layer. The tribe Cynoglosseae has many common characters of pollen grains, but there are some differences among genera. A key to the genera is given based on pollen morphology. Bothriospermum, without endocingulus, may be a primitive genus in this tribe, and its pollen morphology is more similar to that of Eritrichieae than to Cynoglosseae, thus it seems more reasonable to put it into Eritrichieae. Solenanthus may be the most advanced genus in the tribe Cynoglosseae for its pollen grains of sub-isopolar and ovoid shape.