Petrocodon retroflexus sp. nov. (Gesneriaceae) from a karst cave in Guizhou,China

Petrocodon retroflexus Q. Zhang & J. Guo, a new species of Gesneriaceae from Guizhou, China, is described and illustrated. The new species is morphologically similar to Petrocodon viridescens W. H. Chen, Mich. Möller & Y. M. Shui, but differs by its leaf blade being ovate, broadly ovate to elliptic, 8–20 × 5–15 cm and densely pilose on both surfaces, narrowly triangular, 9 mm long, densely pilose bracts, white corolla that is decurved near the middle, with adaxial lip shallowly 4‐partitioned and retroflexed, and pistil ca 1.3 cm long and densely pilose and glandular‐pilose.     

Oreocharis duyunensis (Gesneriaceae), a new species from Guizhou,China

Oreocharis duyunensis Z.Y. Li, X.G. Xiang et Z.Y. Guo (Gesneriaceae), endemic to Guizhou, China, is described as a new species. The new species is easily distinguished from other Oreocharis species by having a white, stellate ring‐like disc. Morphologically, the species is similar to Oreocharis chienii (Chun) Mich. Möller & A. Weber but differs by its densely white pilose leaves, mixed brown‐pilose, pubescent and glandular‐pubescent peduncle and pedicle, white stellate ring‐like disc, and linear‐lanceolate capsule. Based on phylogenetic reconstruction and interspecific genetic distances of nuclear ITS and chloroplast trnL‐F regions, O. duyunensis is a new species with uncertain phylogenetic position.     

PRODUCTION OF HYALINE HAIRS BY INTERTIDAL SPECIES OF FUCUS (FUCALES) AND THEIR ROLE IN PHOSPHATE UPTAKE1

A field study to determine the precise times of year at which three intertidal species of Fucus start to produce hyaline hairs and cease producing such hairs was conducted on the Isle of Man, U.K. Hairs were first observed during February, and within 6 days of their initial appearance, all tagged plants of all species at all tidal heights on the shore possessed hairs. Hair production continued until the beginning of October, at which time Fucus plants growing at the lowest stations (+ 3.0 m) had glabrous apical growth. Hair production continued later into the year for plants growing higher on the shore, and it was not until mid-November that glabrous apical growth was observed in all plants. Phosphate uptake rates of pilose (hairy) and glabrous (hairless) apical sections were measured in November 1988 for F. spiralis L. and in January 1989 for F. spiralis and F. serratus L., at phosphate concentrations ranging from 0.8 μM (ambient seawater) to 9.0 μM. In ambient seawater, pilose plants of F. spiralis removed phosphate 2–3 times faster than glabrous plants, whereas the uptake rates of pilose plants of F. serratus were about 50% greater than those of glabrous plants. The differences between uptake rates of pilose and glabrous plants of both species were smaller or nonsignificant at higher phosphate concentrations. The field and laboratory data are consistent with the hypothesis that hairs are formed in Fucus as a response to increased nutrient demand and that hairs facilitate the uptake of nutrients from seawater at concentrations typical of natural situations.     

Comparative morphology of the leaf epidermis in the genera Codonanthe (Martius) Hanstein and Nematanthus Schrader (Gesneriaceae)*

The leaf epidermis of 14 species ofCodonanthe and 10 species of Nematanthus has been examined. Species of Codonanthe section Codonanthe are geographically restricted to south-eastern Brazil, and are diploid. They possess multicellular-uniseriate nonglandular trichomes, glandular trichomes with a four-celled head and a short body, anisocytic stomata and lack extrafloral nectaries. Species of Codonanthe section Spathuliformae and Codonanthe subgenus Codonanthella are distributed from southern Mexico through Central America to north-western South America and are tetraploid. They possess unicellular non-glandular trichomes (except C. caribaea), glandular trichomes with a two-celled head (except C. caribaea) and a short body, anisocytic stomata and extrafloral nectaries (except C. caribaea). All Nematanthus species are distributed in south-eastern Brazil and are diploid (N=8). Six species of Nematanthus consistently have multicellular-uniseriate nonglandular trichomes, glandular trichomes with a four-celled head and a short (unicellular) or long (multicellular) body, anisocytic stomata and lack extrafloral nectaries. Four species of Nematanthus have multicellular-uniseriate non-glandular trichomes, glandular trichomes with a head of more than four cells and a short body, anisocytic and helicocytic stomata and lack extrafloral nectaries.     

Trichome micromorphology of Iranian Stachys (Lamiaceae) with emphasis on its systematic implication

Trichomes of 37 taxa of the genus Stachys and one species of Sideritis (S. montana) were examined using light and scanning electron microscopy. The indumentum shows considerable variability among different species, but is constant among different populations of one species, and therefore, affords valuable characters in delimitation of sections and species. The characters of taxonomic interest were presence of glandular and non-glandular trichomes, thickness of the cell walls, number of cells (unicellular or multi-cellular), presence of branched (dendroid) trichomes, presence of vermiform trichomes, orientation of trichomes in relation to the epidermal surface, curviness of trichomes, and presence of papillae on trichome surface. Two basic types of trichomes can be distinguished: glandular and non-glandular trichomes. The glandular trichomes can in turn be subdivided into subtypes: stalked, subsessile, or sessile. The stalks of the glandular trichomes can be uni- or multi-cellular. Simple unbranched and branched trichomes constitute two subtypes of non-glandular trichomes. Our data do not provide any support for separation of Sideritis from Stachys. The following evolutionary trends are suggested here for Stachys: vermiform trichomes with stellate base are primitive against vermiform trichomes with tuberculate base, long vermiform trichomes are primitive against the short simple trichomes, appressed trichomes are advanced against spreading ones, and loss of glandular trichomes is advanced against their presence. Overall, trichome micromorphology is more useful in separation of species within sections rather than characterizing large natural groups known as sections, except for few cases.     

No benefit of glandular trichome production in natural populations of Datura wrightii?

Populations of Datura wrightii vary in the frequency of plants that produce glandular trichomes, a resistance trait under the control of a single gene. Such variation may be maintained if the production of glandular trichomes is costly in the absence of herbivory, and if selection imposed by herbivore communities varies spatially or temporally. Here, we document costs in the presence of herbivory for established glandular plants relative to established non-glandular plants growing in natural populations from coastal mountain, Riversidian sage scrub, and Mojave desert habitats in southern California. Damage caused by the herbivore community varied spatially, with significant differences in herbivore-specific damage between plants of the two trichome types and among populations within habitats, although not generally among habitats. Plants with greater canopy size and canopy persistence had higher viable seed production than smaller or more damaged plants, but this relationship was statistically significant only for non-glandular plants. However, the relationship between viable seed production and canopy persistence became significant for glandular plants when damage caused by sap suckers, which do not remove leaf area, was pooled with undamaged leaf area. The high cost exhibited by glandular plants leads us to predict that in the absence of any additional, unknown benefits of producing glandular trichomes, the frequency of these plants should decline in all natural populations of D. wrightii. Received: 13 July 1999 / Accepted: 28 September 1999     

Indirect cost of a defensive trait: variation in trichome type affects the natural enemies of herbivorous insects on <Emphasis Type="Italic">Datura wrightii</Emphasis>

The costs and benefits of defensive traits in plants can have an ecological component that arises from the effect of defenses on the natural enemies of herbivores. We tested if glandular trichomes in Datura wrightii, a trait that confers resistance to several species of herbivorous insects, impose an ecological cost by decreasing rates of predation by the natural enemies of herbivores. For two common herbivores of D. wrightii, Lema daturaphila and Tupiocoris notatus, several generalized species of natural enemies exhibited lower rates of predation on glandular compared to non-glandular plants. Lower rates of predation were associated with reductions in the residence time and foraging efficiency of natural enemies on plants with glandular trichomes, but not with direct toxic effects of glandular exudate. Our results suggest that the benefit of resistance to herbivores conferred by glandular trichomes might be offset by the detrimental effect of this trait on the natural enemies of herbivores, and that the fitness consequences of this trichome defense might depend on the composition and abundance of the natural-enemy community.     

Sound communication in social voles (subgenus Sumeriomys)

ABSTRACT

The acoustic communication of three species of social voles from the subgenus Sumeriomys – Microtus socialis (two subspecies: M. s. socialis and M. s. goriensis), M. paradoxus and M. hartingi – are described. Vole sound communication includes two main signals: squeaks and singing. The sounds made by M. hartingi have significantly higher frequency parameters than those of other species. Voles of all species squeak in situations of distress, and the males sing during courtship of the females. However, singing in social voles is not a necessary pattern for sexual behaviour: less than half of M. s. socialis and M. paradoxus males sang, M. hartingi sang even more rarely and M. s. goriensis did not demonstrate this behaviour at all. Despite the great similarity of the squeaks, its parameters differ significantly between species and differ from those of the common voles. This introduces one more argument that M. paradoxus and M. socialis are independent species, as are the subgenera Sumeriomys and Microtus.     

Micromorphological and chemical characterisation of Stachys recta L. subsp. serpentini (Fiori) Arrigoni in comparison to Stachys recta L. subsp. recta (Lamiaceae)

Stachys recta L. is a very polymorphous species in which numerous subspecies were recognised. S. recta L. subsp. serpentini (Fiori) Arrigoni is a typical endemism growing on serpentine soils in northern Apennines and particularly in Tuscany (Italy). In order to contribute to a better knowledge of this plant and to its differentiation with respect to S. recta L. subsp. recta, the micromorphological characters (non-glandular and glandular trichomes) and the essential oil composition of the two subspecies were investigated. Micromorphological characters were studied using scanning and transmission electron microscopy, while light microscopy was used for histochemical observations. Essential oil analysis was carried out by gas chromatography and mass spectrometry.In the two examined taxa, the morphology and distribution of glandular and non-glandular trichomes, and the different essential oil composition, may be considered distinctive characters at subspecies level. This is consistent with the taxonomic classification considering S. recta subsp. serpentini a subordinate taxon of S. recta.     

Seed predation and dispersal of glabrous filbert (Corylus Heterophylla) and pilose filbert (Corylus Mandshurica) by small mammals in a temperate forest, northeast China

We investigated the seed dispersal of glabrous filbert (Corylus heterophylla) and pilose filbert (Corylus mandshurica), two large-seeded shrub species in a temperate forest, northeast China, September 2006. Small mammals such as Apodemus speciosus, Clethrlonomys rufocanus, and Eutamias sibiricus, were regarded as the main dispersal agents. More seeds were harvested by small mammals in pilose filbert (98%) than in glabrous filbert (87.5%) till our last survey. Seed removal rates differed between the two species. Fewer seeds of glabrous filbert (17.5%) were eaten in situ than pilose filbert (57.5%). More seeds of glabrous filbert were removed (70%), stay intact after removal (25.5%), eaten after removal (16%) than pilose filbert. However, more seeds were cached after removal in pilose filbert than in glabrous filbert (10.5 and 4%, respectively). Fewer tagged seeds of pilose filberts (14%) were missed than glabrous filberts (24.5%). About 8 and 12 primary caches were found in glabrous filbert and pilose filbert seeds respectively, indicating scatter hoarding. All of the removed seeds were distributed within 10 m of seed stations for both filberts. The average dispersal distances for glabrous filbert did not differ from pilose filbert. Only a small proportion of the caches remained till our last survey (2 and 1%, respectively). Based on the results, we found a difference in dispersal patterns of glabrous filbert and pilose filbert seeds. Evidences showed that glabrous filberts might be a less preferred seed species for small seed-eating mammals compared with pilose filbert, probably due to its harder and thicker husk and low seed profitability.     

Morphological and anatomical analyses of rheophytic Rhododendron ripense Makino (Ericaceae)

The comparative morphology and anatomy of the leaves of the rheophytic Rhododendron ripense and the closely related inland species Rhododendron macrosepalum were examined. The leaf of R. ripense is thinner than that of R. macrosepalum, with leaf length to width ratios (leaf index) of 2.92 and 1.91, respectively. Moreover, the leaf of R. ripense consists of fewer cells than the leaf of R. macrosepalum, suggesting stenophyllization of R. ripense caused by the decreased number of cells. In addition, leaf thickness and the number of stomata per leaf of R. ripense were significantly greater than those of R. macrosepalum, but the density of the short glandular pilose hairs on the leaf of R. ripense was lower. The observed morphological differences between the two species may be explained by certain aspects of the riparian environment, such as high irradiation and frequent flooding after heavy rainfall, to which R. ripense is exposed.     

The effect of flowering calendula and cuphea plants on Orius insidiosus survival and predation of Aphis glycines

Flowering oilseed crops have the potential to diversify agroecosystems currently dominated by corn and soybeans and improve the provision of ecosystem services such as pest control. Nectar and pollen feeding may increase natural enemy fitness and searching behaviour, increasing their survival and prey consumption rates. The soybean aphid (Aphis glycines Matsumura; Hemiptera: Aphididae) is a particularly widespread and costly agricultural pest. In this study, we evaluate the effects of two flowering oilseed crops, cuphea and calendula, on the survival of the insidious flower bug (Orius insidiosus Say; Hemiptera: Anthocoridae) and its consumption levels of A. glycines placed on soybean plants. We also evaluated the survival of O. insidiosus when placed on glandular and non-glandular cuphea varieties. The amount of A. glycines that remained unconsumed by O. insidiosus did not differ among treatments. Because mortality levels of O. insidiosus were higher on glandular compared to non-glandular cuphea plants, glandular trichomes, or plant hairs, may play a role in impeding movement and prey consumption by O. insidious.     

国产柿属(Diospyros)的叶表皮形态研究及其分类学意义

为了解柿属(Diospyros)植物的叶表皮特征,在光学显微镜下观察了46种2变种植物的叶表皮微形态特征,并进行了主成分分析。结果表明,柿属植物的叶表皮微形态可分为2组：A组的表皮细胞为不规则形,垂周壁浅波状或深波状;B组的表皮细胞为多边形,垂周壁平直或弓形。气孔全部位于下表皮,以无规则型为主,但同时有辐射型、环列型和十字型气孔存在。大多数种具有表皮毛,表皮毛有非腺毛和头状腺毛的区别。柿属植物的叶表皮微形态特征类型多样,能够为部分类群的分类提供较好的形态学证据。     

DIFFERENTIAL EFFECTS OF PHOTOPHASE IRRADIANCE ON METABOLIC AND URINARY STRESS HORMONE CONCENTRATIONS IN BLIND AND SIGHTED RODENTS

The effects of different photophase irradiance levels on the daily rhythms of energy expenditure (DEE, calculated from oxygen consumption, VO₂) and urinary metabolites of stress hormones in sighted (Microtus socialis) and blind (Spalax ehrenbergi) rodents were compared. Five groups of each species were exposed to different irradiance levels (73, 147, 293, 366, and 498 μW/cm²) under short photoperiod (8L:16D) condition with constant ambient temperature 25?±?2°C for 21 days before assessments. As light intensity increased from 73 μW/cm², both species reduced DEE, especially among M. socialis. Cosinor analysis revealed significant ultradian rhythms in VO₂ of M. socialis with period length being inversely related to irradiance level. Conversely, in S. ehrenbergi, robust 24 h VO₂ rhythms were detected at all irradiances. In M. socialis, significant 24 h rhythms in urinary output of adrenaline were detected only at 293 μW/cm², whereas for cortisol, unambiguous rhythms were detected at 73 and 147 μW/cm². Distinct adrenaline daily rhythms of S. ehrenbergi were observed at 73 and 293 μW/cm², whereas this species exhibited significant rhythms in cortisol at 147 and 293 μW/cm². Changes in photophase irradiance levels affected stress hormone concentrations in a dose-dependent manner. There were significant negative and positive correlations of M. socialis and S. ehrenbergi stress hormones, respectively, with increasing irradiance. Our results indicate photophase light intensity is another environmental factor that can significantly affect entrainment of mammalian daily rhythms. Both low and high irradiance conditions can trigger stress responses, depending on the species' natural habitat. (Author correspondence: zubidat3@013.net.il)     

A Study on the Morphology of Foliar Trichomes of Evergreen Oaks (Quercus) in China and Its Implication in Taxonomy

Most of the evergreen oaks (Quercus L.) are endemic to China and distributed in a large moutainous area of southwestern China at an altitude of 2600-4000m. The delimitation of sections and species in the group has not been satisfactorily solved. The foliar trichomes are very an important character in identifying species of oaks. As a result of observation on 17 species and 2 varieties of evergreen oaks in China under scanning electron microscope, ten types of foliar trichomes are recognized: Simpleuniseriate, Simple-branched, Bulbous, Rosulate, Solitary, Stipitate-fasciculate, Fasciculate, Multiradiate, Stellate and Fused-stellate. The first four types fall into glandular trichomes and the last six non-glandular trichomes. The taxa examined have a combination of various types of the trichomes. All the evergreen oaks have non-glandular trichomes on their foliar epidermis, but glandular trichomes occur solely in certain taxa. These two types of foliar trichomes are obviously different in structure and function, which represent different adular trichomes and the last six non-glandular en oaks are divided into two groups: the glandular group and the non-glandular group according to the types of foliar trichomes and it is reasonable to divide the evergreen oaks into two sections: Sect. Suber (Reichenb.) Spach and Sect. Englerianae (A. Camus) Hsu et Jen. Foliar trichomes of the evergreen oaks show a continuity in density from species predominately with glandular trichomes at a high altitude to species solely with non-glandular trichomes at a low elevation. The characters of branching of foliar trichomes, especially the number of branches, can imply to some extent the evolutionary position of a given evergreen species. Considering the types, density and branches of foliar trichomes combined the evolutionary relationships among the evergreen oaks in China are proposed. Q. gilliana is the startpoint, from where evolution took place in two directions: glandular and non-glandular. In the glandular direction envolved are Q. aquifolioides, Q. longispica and Q. guyavaefolia. Q. pannosa is a species with the most types and highest number of glandular trichomes. In non-glandular line aligned are Q. spinosa, Q. senescens, Q. senescens var. muliensis, Q. monimotricha with the reduction of glandular trichomes and increase of non-glandular trichomes. After Q. tungmaiensis no glandular trichomes appear and it solely depends on non-glandular trichomes to protect itself. Q. granchetii is the climax in the non-glandular line with the highest density and number of branches of foliar trichomes. And then some species, such as Q. engleriana, Q. phillyraeoides, have few or no trichomes. Q. dolicholepis, Q. baronii and Q. baronii var. capitata show the close re-lationships because they share fused-stellate trichomes.     

Diverging patterns of host use by phytophagous insects in relation to leaf pubescence in Arbutus xalapensis (Ericaceae)

Summary Foliar pubescence in the mexican madrone (Arbutus xalapensis H.B.K.) is an extremely variable character. Leaf specimens of pilose and glabrous madrones showed a similar composition of major secondary compounds. On the other hand, sympatric pilose and glabrous individuals were found to support significantly different insect guilds. Insect preference was strongly associated to mouthpart anatomy. Chewing and gall-forming insects were significantly more abundant on glabrous trees while sucking insects were more common on pilose trees. Additionally, parasitation of sucking insects was lower on pilose trees, possibly because the plant trichomes provide protection against parasitoids.     

Nutlet morphology in the tribe Elsholtzieae (Lamiaceae)

A comprehensive study on nutlet morphology on 46 taxa from the tribe Elsholtzieae was conducted using stereomicroscopy, light microscopy and scanning electron microscopy. The objectives were to evaluate the nutlet characteristics in a systematics context and to provide additional evidence for taxon delimitation. The nutlet shape varied from elliptical, widely elliptical, spheroidal and ovate to pyriform. The size ranged from 0.50 to 2.70 mm in length and 0.34 to 2.65 mm in width. The three types of trichomes included simple non-glandular trichomes (Collinsonia punctata), clavate non-glandular trichomes (Elsholtzia cephalantha) and peltate glandular trichomes (Mosla grosseserrata and M. chinensis). We also report the presence of a mucilaginous reaction in nine species of the genus Elsholtzia in Elsholtzieae for the first time. Four types (undulate, papillae-like projection, thorn-like pattern and reticulate) and three subtypes (reticulate, reticulate-cellular and supra-reticulate) of nutlet surface patterns in the mid-ventral region were recognized in Elsholtzieae based on the differences in cell outline, outer periclinal walls and dentations of the anticlinal walls. These nutlet micromorphological results may serve as a reference for future hypotheses on the classification of the tribe Elsholtzieae.     

Trichoderma species form endophytic associations within Theobroma cacao trichomes

Trichoderma species are usually considered soil organisms that colonize plant roots, sometimes forming a symbiotic relationship. Recent studies demonstrate that Trichoderma species are also capable of colonizing the above ground tissues of Theobroma cacao (cacao) in what has been characterized as an endophytic relationship. Trichoderma species can be re-isolated from surface sterilized cacao stem tissue, including the bark and xylem, the apical meristem, and to a lesser degree from leaves. SEM analysis of cacao stems colonized by strains of four Trichoderma species (Trichoderma ovalisporum-DIS 70a, Trichoderma hamatum-DIS 219b, Trichoderma koningiopsis-DIS 172ai, or Trichoderma harzianum-DIS 219f) showed a preference for surface colonization of glandular trichomes versus non-glandular trichomes. The Trichoderma strains colonized the glandular trichome tips and formed swellings resembling appresoria. Hyphae were observed emerging from the glandular trichomes on surface sterilized stems from cacao seedlings that had been inoculated with each of the four Trichoderma strains. Fungal hyphae were observed under the microscope emerging from the trichomes as soon as 6 h after their isolation from surface sterilized cacao seedling stems. Hyphae were also observed, in some cases, emerging from stalk cells opposite the trichome head. Repeated single trichome/hyphae isolations verified that the emerging hyphae were the Trichoderma strains with which the cacao seedlings had been inoculated. Strains of four Trichoderma species were able to enter glandular trichomes during the colonization of cacao stems where they survived surface sterilization and could be re-isolated. The penetration of cacao trichomes may provide the entry point for Trichoderma species into the cacao stem allowing systemic colonization of this tissue.     

CALCIFICATION IN THE GREEN ALGA HALIMEDA. I. AN ULTRASTRUCTURE STUDY OF THALLUS DEVELOPMENT1

The ultrastructure of 4 species of the calcareous, siphonaceous alga Halimeda (H. cylindracea Decaisne, H. discoidea Decaisne, H. macroloba Decaisne and H. tuna (Ellis & Solander) Lamour) has been studied, and the observed changes during growth and development are related to changes in the degree of calcification. A distinct gradient in the types and quantities of cell organelles exists in a growing apical filament. As these filaments grow, branch, and eventually develop into a mature segment, changes in the organization of organelles such as mitochondria and chloroplasts are observed. Calcification begins when the chloroplasts reach structural maturity and when the peripheral utricles adhere (fuse). This adhesion of the peripheral utricles isolates the intercellular space (ICS) in which calcification occurs from the external seawater. Calcification begins in the outermost (pilose) cell wall layer of the walls facing into the ICS. The cell walls at the thallus exterior undergo extensive changes after utricular fusion; the pilose layer is lost, the cuticles of adjacent utricles fuse forming a ridge at their junction, and multiple cuticles are formed. The aragonite (CaCO₃) crystals which are initially precipitated within the pilose wall layer, rapidly increase in size and number, eventually filling much of the ICS. Only the initial nucleation of aragonite is associated with the pilose wall layer, the later precipitation of aragonite is totally independent of the pilose layer. In older segments secondary deposition of CaCO₃ also occurs around existing aragonite needles.     

The anatomical and micromorphological characteristics of <Emphasis Type="Italic">Salvia vermifolia</Emphasis> (Section <Emphasis Type="Italic">Aethiopis</Emphasis> Bentham,Lamiaceae) from Central Anatolia,Turkey

The gross anatomical features, namely stem, blade and petiole and the micromorphological features, trichome, pollen and nutlet of Salvia vermifolia Hedge & Hub.-Mor. (Sect. Aethiopis Bentham, Lamiaceae) were examined using light microscopy (LM) and scanning electron microscopy (SEM). S. vermifolia is a perennial endemic herb growing on igneous and serpentine slopes in Sivas province, Central Anatolia, Turkey. The results obtained from anatomical studies show that the stem is made up of 4–8-layers of collenchyma cells and 1-3-layers chlorenchyma cells. The blade is amphistomatic, bifacial, with 2-3-layered palisade cells on the adaxial. The petiole has three large central bundles and six small subsidiary bundles. Peltate glandular, capitate glandular and non-glandular trichomes are present in S. vermifolia. The peltate glandular trichomes are present in abundance on the calyx and corolla, and capitate glandular trichomes are predominant on the calyx, inflorescence axis, pedicel and stem. The acicular non-glandular trichomes are mainly situated on the calyx, corolla, inflorescence axis and pedicels whereas the floccose non-glandular trichomes are common on both sides of the leaf, inflorescence axis and petiole. The pollen grains are hexacolpate to octacolpate, medium to large (P, polar axis=45.29–62.49 μm; E, equatorial axis=45.32-62.38 μm), oblate-spheroidal to prolate-spheroidal (P/E=0.91–1.02) and their exine ornamentation is bireticulate. The nutlets are 2.70–2.90 mm long and 1.90–2.13 mm wide, brown, ovoid-oblong in their outline, rounded-trigonous in transverse section, and their surface is glabrous and colliculate.