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 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 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 THE RELATIONSHIPS OF SIMMONDSIA CHINENSIS TO THE ORDER EUPHORBIALES

Pollen from Simmondsia chinensis (Simmondsiaceae) was examined in LM, SEM, and TEM. The pollen is shed as monads, triangular in shape in polar view, with a 3-porate aperture type in which the pores are large and poorly defined. The tectum is irregularly scabrate, sometimes forming minute “islands” topped with spinules. In thin section, the endexine is thickened and lamellate in the aperture regions, and narrow in the mesoporus; the foot layer is well-defined but noticeably thicker in the mesoporus; and thin columellae support an essentially complete tectum. The pollen of four genera, Buxus, Pachysandra, Sarcococca, and Styloceras, from the Buxaceae to which Simmondsia has been assigned by some authors, was also examined and illustrated. The pollen morphology of two families frequently aligned with Simmondsiaceae, Euphorbiaceae and Pandaceae, is briefly discussed. For the most part pollen morphology supports the treatment of Simmondsia as a monotypic family, Simmondsiaceae.     

PALYNOLOGY AND SYSTEMATICS OF CUPHEA (LYTHRACEAE). I. MORPHOLOGY AND ULTRASTRUCTURE OF THE POLLEN WALL

A survey of pollen from 153 species of Cuphea has revealed a remarkable array of morphological forms. The survey involved light and electron microscope investigations of C. crassiflora, jorullensis, and koehneana to determine details of exine morphology, and a more general study of pollen from an additional 150 species. Comparison of pollen types within a single morphological category and within taxonomic groups (i.e., sections or subsections) indicates considerable variation at subgeneric levels. The genus is distinctly eurypalynous, and the extent to which pollen varies among the sections, subsections, species and varieties is probably exceeded by few genera of comparable size. The pollen is also variable within each taxon, but size studies of single-anther lactic acid preparations demonstrate the genus is not polymorphic, as in Lythrum, where pollen polymorphism is associated with heterostyly. The latter phenomenon is as yet unknown in Cuphea. These results reveal that pollen morphology constitutes an important and useful character for taxonomic studies of Cuphea.     

THE POLLINATION ECOLOGY OF SOLANUM ROSTRATUM (SOLANACEAE)

The pollination mechanisms and pollen vectors of Solarium rostratum have been examined by greenhouse experiments and field studies. Although the capacity for autogamy exists in this weedy annual, it rarely occurs because of two factors: (1) the morphology of the flower and (2) the foraging behavior of the various species of Bombus, the primary pollen vector in the regions studied. The percentages of geitonogamy and xenogamy are dependent on the flight pattern of the bees and the number of open flowers on a plant.     

FUCHSIA POLLEN FROM THE EARLY MIOCENE OF NEW ZEALAND

Fossil pollen of Diporites aspis Pocknall & Mildenhall, from the early Miocene of New Zealand, was examined by combined light, scanning, and transmission electron microscopy, in order to determine its relationship to extant Fuchsia pollen. Based on overall morphology, exine structure, exine surface sculpture, viscin thread morphology, and apertural features, the fossil pollen definitely can be considered to represent Fuchsia. Parallel studies of extant Fuchsia pollen indicate that a more precise identification of the fossils is not possible. With the examination of additional fossil material, it should be possible to learn a great deal about the timing of origin and migration of the genus Fuchsia, but not of its constituent parts.     

EVOLUTION OF EXINE STRUCTURE IN THE POLLEN OF PRIMITIVE ANGIOSPERMS

Diversity in the structure of the exine in 35 families of the ranalean complex is compared through a series of representative scanning electronmicrographs, and evolutionary trends in exine structure of primitive angiosperms are outlined, along with discussion of the significance of these data for understanding the evolution of exine structure in flowering plants as a whole. In order to reduce ambiguity in the palynological literature, it is suggested that persons undertaking light microscope studies of unstained, acetolyzed pollen grains adopt the morphological terms sexine-nexine in describing pollen wall layers while restricting their use of the chemically defined terms ektexine-endexine largely to pollen studies carried out with the transmission electron microscope. This study emphasizes that a clear understanding of the palynological concept of structure versus sculpturing is a necessary prerequisite for the taxonomic/ phylogenetic use of pollen wall morphology. Finally, data from investigation of a number of ranalean families of primitive angiosperms support the conclusion that the direction of a recurrent and major evolutionary trend in exine structure of flowering plants proceeds from pollen that is tectate-imperforate to tectate-perforate pollen to semitectate pollen, and more rarely, to pollen grains that are intectate.     

THE AURICULATE POLLEN GRAIN OF HYMENOCALLIS QUITOENSIS HERB. (AMARYLLIDACEAE) AND ITS SYSTEMATIC IMPLICATIONS

The pollen grain of Hymenocallis quitoensis Herb, is examined with scanning and transmission electron microscopy. The very large, monosulcate grain is characterized by a coarsely reticulate exine surface and solid, auriculate appendages at the equatorial facies. Surface morphology of the pollen grain of H. quitoensis is compared with other representative species of Hymenocallis and supports the inclusion of H. quitoensis within the genus. Meridional exine dimorphism in other species of Hymenocallis is interpreted as a reduction from the well-defined auriculae of H. quitoensis. Surface morphology of the pollen grain of Hymenocallis sects. Artema Traub and Elisena (Herb.) Traub exhibits divergence from that of sects. Hymenocallis and Ismene (Herb.) Traub. Phylogenetic and paleobotanical implications of the auriculate pollen grain are briefly discussed.     

POLLEN MORPHOLOGY OF THE DILLENIACEAE AND ACTINIDIACEAE

Pollen of 53 species of Dilleniaceae and Actinidiaceae was examined by light microscopy, scanning electron microscopy, and a selected group in transmission electron microscopy. Dilleniaceae pollen ranges from tricolpate, tricolporate, tetracolpate, and incipiently inaperturate. Tricolpate types occur only among the Old World subfamily Dillenioideae and the compound aperturate (3-colporate) condition is restricted to the subfamily Tetraceroideae. Within the Dilleniaceae the tricolpate pollen type with elongated apertures is considered primitive, having given rise to the 3-colporate and 4-colpate conditions. The striking pollen dimorphism in the Neotropical species of Tetracera, all of which are androdioecious, is documented; however, in contrast to previous reports, pollen from bisexual flowers appears to be incipiently inaperturate and not pantoporate. The inaperturate condition is interpretated as an early stage in the evolution of outcrossing. Pollen morphology does not support a close relationship between Dilleniaceae and Actinidiaceae. Pollen morphological differences that can be noted between these families are: tectum complete and predominantly psilate or psilate-granular in Actinidiaceae, tectum incomplete, punctate to reticulate in Dilleniaceae; an equatorial bridge of ektexine over the endoaperture usually present in Actinidiaceae, absent in Dilleniaceae; columellae reduced in Actinidiaceae, columellae usually well-developed in Dilleniaceae. Pollen morphology does not argue against a close relationship between Actinidiaceae and Theaceae.     

CONTRIBUTION OF POLLEN MORPHOLOGY TO SYSTEMATICS OF COLLOMIA (POLEMONIACEAE)

Pollen morphology of 14 species of Collomia (Polemoniaceae) was examined by light microscopy, and by both scanning and transmission electron microscopy. Four distinct pollen types were observed which are based principally upon 1) shape, number and distribution of apertures, and 2) surface sculpturing: Type 1—zonocolporate with striate ridges; Type 2—zonocolporate with striato-reticulate ridges; Type 3—pantoporate with radiate ridges; Type 4—pantoporate with irregularly reticulate ridges. Evaluation of pollen morphology reveals considerable discrepancy with respect to presently accepted sectional classification. Collomia grandiflora of sect. Collomia has a pollen type similar to that of members of sect. Collomiastrum and is now interpreted as representing an independent evolutionary line derived from the latter section. Collomia diversifolia of sect. Courtoisia has a pollen morphology similar to that of sect. Collomia. whereas C. heterophylla of the same section possesses pollen unique within the genus. This last pollen type shows close similarity to the pollen of members of Polemonium, Gilia, Leptodactylon, and Ipomopsis. Pollen of C. tinctoria and C. tracyi of sect. Collomia are anomalous within Polemoniaceae. No significant difference in exine stratification was discernible among the four pollen types.     

中国马鞭草科植物的花粉形态

本文对我国18属91种3变种马鞭草科(Verbenaceae)植物的花粉形态进行了详细地观察研究。其中对15属27种进行了扫描电镜观察。本科为花粉多类型的科之一。根据萌发孔的数目、特征和纹饰特点,本文将马鞭草科分为9个花粉类型:1.具疣3沟花粉类型;2.具刺3沟花粉类型;3.具细网8沟花粉类型;4.具模糊颗粒-细网3沟花粉类型;5.具网3沟花粉类型;6.具散沟(6-8条)花粉类型;7.具3孔花粉类型;3.具细网3沟花粉类型,4.具模糊颗粒一细网3沟花粉类型,5.具网3沟花粉类型,6.具散沟(6-8条)花粉类型,7.具3孔花粉类型,8.具3孔沟花粉类型,9.具3(4)孔沟花粉类型。     

POLLEN OF BOERLAGIODENDRON: A UNIQUE TYPE IN THE ARALIACEAE

The pollen of Boerlagiodendron, one of the most distinct genera of Araliaceae, is unique within the family in its remarkable degree of differentiation of sexine into pertectate and intectate patterns and in its reduced apertures. The unexpanded mesocolpia and the inconspicuous apertures have apparently caused misinterpretation of the pollen morphology of B. pectinatum in recent literature. A comparative study of several pollen features, together with the meristic characters of carpels and stamens, has demonstrated some significant correlations among them. It appears that the origin of the genus may have been centered around the New Guinea and Solomons areas.     

THE PLATYCARYA PERPLEX AND THE EVOLUTION OF THE JUGLANDACEAE

We report on the leaves, fruits, inflorescences, and pollen of two fossil species in the genus Platycarya. The association of these dispersed organs has been established by their repeated co-occurrence at a large number of localities, and for two of the organs (fruit and pistillate inflorescence, and pollen and staminate inflorescence) by apparent organic attachment of compression fossils. Each of the two species can be distinguished by characteristics of all the known megafossil organs. We also review the fossil record of dispersed platycaryoid fruits and inflorescences, recognizing three additional species of Platycarya and two of Hooleya. Two of the fossil Platycarya species are morphologically very different from the living Platycarya strobilacea Sieb. et Zucc., but they show the diagnostic features of the genus. Hooleya is a generalized member of the Platycaryeae that is probably close to the ancestry of Platycarya. The two Platycarya species known from multiple organs provide a remarkable example of mosaic evolution in which fertile and foliar structures have attained different levels of morphological specialization. The leaves, often considered the most plastic of plant organs, retain several features that are otherwise seen only in the Engelhardieae. These similarities in leaf architecture between the fossil Platycarya species and Engelhardieae are advanced features for the Juglandaceae, and thus indicate a sister-group relationship between the two lines. In contrast to the leaves, the fruits, inflorescences, and pollen of the fossil Platycarya species are almost as specialized as those of the extant P. strobilacea and bear little resemblance to the same structures in other genera of the family. The morphology, taphonomy, sedimentary setting, and geographic and stratigraphic distribution of three of the fossil platycaryoid species suggest that they were wind-dispersed, early successional plants that grew in thickets. This habit is retained by Platycarya strobilacea and is typical of many of the amentiferae (e.g. Myricaceae, Betulaceae). The r-selected life-history pattern of the Platycarya line may well have contributed to its low diversity through geologic time.     

THE MOTION OF WINDBORNE POLLEN GRAINS AROUND CONIFER OVULATE CONES: IMPLICATIONS ON WIND POLLINATION

Comparisons are presented between the three-dimensional airflow patterns created around and by a scale model of a conifer ovulate cone and the trajectories of windborne pollen grains around Picea, Larix, and Pinus ovulate cones. Three general components of the airflow pattern around an ovulate cone model are 1) doldrum-like eddies, rotating over the adaxial surfaces of cone scales and directed toward attached ovules, 2) airflow spiralling around the cone axis along cone scale orthostichies and parastichies, and 3) a complex pattern of vortices (“umbilicus”) directed toward the leeward surface of the ovulate cone. The observed trajectories of pollen grains around cones of Picea, Larix, and Pinus conform to two of these three airflow components: 1) pollen grains are seen to roll along cone scales toward the distal scale margin and to become reentrained in airflow directed backward toward attached ovules, and 2) pollen grains passing around the cone are deflected into the “umbilicus” airflow pattern, where they either settle on or impact with cone scales (approach trajectories), or where they approach the leeward cone surface but are deflected away by airflow passing under the cone (Z-shaped trajectories). Vectoral analyses of pollen grain motion reveal a complex pattern of trajectories influenced by boundary layer conditions defined by ovulate cone geometry and ambient airflow speed. Wind tunnel studies of ovulate cones subtended by leaves and stem indicate that leaves circumscribing the cone act as a snowfence, deflecting windborne pollen toward the cone. Vectoral analyses of airflow patterns and pollen grain trajectories close to ovulate cones indicate that wind pollination in conifers is a non-stochastic aerodynamic process influenced by cone-leaf morphology and the behavior of pollen grains as windborne particles.     

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE. XII. THE FLORAL BIOLOGY OF CABOMBA CAROLINIANA

Observations have been made on the pollination ecology of Cabomba caroliniana Gray in Texas. Flowers are trimerous with morphologically similar perianth parts. The adaxial corolla spurs are nectariferous and attract small Diptera (e.g. Notiphila cressoni and Hydrellia bilobifera). Anthesis occurs for 2 consecutive days with flowers opening about 10:00 a.m. and closing around 4 p.m. on each day. First-day flowers have short, indehiscent stamens and longer pollen-receptive stigmata which arch outward over the nectaries. In 2nd-day flowers the stamens have elongated to the level of the stigmata and extrorse dehiscense occurs above the nectaries. Stigmata of 2nd-day flowers are pressed together at the center of the flower and are nonreceptive to pollen. Insects attracted to 2nd-day flowers in search of nectar become dusted with pollen (due to the position and extrorse dehiscence of the anthers) and as insects fly to 1st-day flowers, achieve cross-pollination by virtue of the stigmata position over the nectaries. Seed anatomy is similar to that of other nymphaeaceous genera (i.e., abundant perisperm, little cellular endosperm, a haustorial nucellar “tube,” and a small dicotyledonous embryo). Pollination morphology and comparative xylem anatomy support the segregation of Cabomba from the Nymphaeaceae, sensu stricto. The anatomical correlations between seeds and the myophilous pollination syndrome (found elsewhere in Nymphaeaceae, sensu lato), however, suggest a phyletic relationship.     

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.     

EQUATIONS FOR THE MOTION OF AIRBORNE POLLEN GRAINS NEAR THE OVULATE ORGANS OF WIND-POLLINATED PLANTS

A technique is presented that is capable of predicting the motion of airborne pollen grains and the probability of pollen capture by wind-pollinated plants. Equations for the motion of rigid-walled particles (= pollen grains, spores, or Sephadex beads) in a supporting, compressible fluid (= air) are derived from the first principles of fluid dynamics. These equations are incorporated into a computer program (MODEL) which can be used with a desktop computer. The operation of MODEL requires empirical data on the pattern of airflow or the motion of a pollen species around the surfaces of the taxonomically relevant ovulate plant organ. With this information, MODEL can predict the behavior of any pollen species for which physical properties (size and density) are specified or empirically known. The significance of this procedure lies in the quantification of physical phenomena that influence the mechanics and fluid dynamics of pollen capture in wind pollination. The technique is illustrated and tested by its application to two grass species (Setaria geniculata and Agrostis hiemalis) for which velocity fields of pollen motion have been previously reported.     

THE CHROMOSOME RACES OF ZINNIA JUNIPERIFOLIA

A comparative study of the distribution, habitat, morphology and cytology of diploid (n = 10) and tetraploid races of Z. juniperifolia of the subgenus Diplothrix (COMPOSITAE-Heliantheae) was carried out to elucidate their relationships. There is a positive correlation between chromosome number and many floral and vegetative characters, including pollen size. Chiasmata frequency is about 0.63 and 0.62 per chromosome in the diploid and in the tetraploid respectively. Multivalents are common in the latter. Although attempts to obtain colchicine-induced tetraploids have failed, available morphological, cytological and chromatographic evidence indicate the tetraploid may be of autoploid origin. Alternative ancestries involving other species of the subgenus are discussed. Taxonomically the tetraploid is considered a race of Z. juniperifolia.     

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.