VARIATIONS IN POLYMNIA POLLEN

Photomicrographs of normal and aberrant Polymnia (Compositae) pollen were made with the scanning electron microscope and with the light microscope. Anomalous pollen forms from F₁ hybrids of P. maculata Cav. are compared to the parental type and to the closely related P. uvedalia (L.) L. Among the variations discussed are differences in size and form, fused aggregates, and reduction in length of spines. The cause of these variations is unknown, although similar anomalies were later found in a single P. maculata parent, Wells No. 740, which grew under similar standard greenhouse conditions.     

POLLEN ORGANS OF THE PENNSYLVANIAN CALLISTOPHYTACEAE (PTERIDOSPERMOPSIDA)

Numerous specimens of the pteridospermous pollen organs Idanothekion glandulosum and Callandrium callistophytoides are present in Middle and Upper Pennsylvanian petrifaction material, respectively. Several specimens of Idanothekion, previously known only in an isolated condition, are attached to foliage assignable to the monostelic seed fern Callistophyton. Proof of the affinities of Idanothekion allows this genus to be elevated from a form genus to an organ genus in the Callistophytaceae. The structure of the foliage and the mode of attachment of the Idanothekion specimens are like those previously described for Callandrium. Structural features of Idanothekion and Callandrium are compared, and the question of their identity is discussed. At present one basic type of pollen organ, borne on the abaxial surface of foliar pinnules, is known for members of the Callistophytaceae.     

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.     

COMPARATIVE POLLEN MORPHOLOGY AND TAXONOMIC AFFINITIES IN CYCADALES

Scanning and transmission electron microscopy of pollen grains of 29 species, representing the ten extant genera of Cycadales, has provided valuable insight into their relationships. Pollen grains of these taxa are boat-shaped, monosulcate, and bilaterally symmetrical. They range from narrowly to widely elliptical or subcircular when viewed distally, and have an exine surface of psilate, foveolate, or fossulate. Pollen wall ultrastructure of Cycadales is typically tectate with alveolate∗∗∗spongy exine. The nexine is laminated in all genera. Nexine 1 (footlayer) is present in most species as a thin and often discontinuous layer. There is consistent variation in thickness of the sporoderm layers among the genera but relative uniformity within them. Pollen characteristics are well correlated with macro- and micromorphological features, chromosome numbers, geographical distribution, and postulated pollination mode. A close affinity between Encephalartos, Lepidozamia, and Macrozamia is recognized. Pollen characteristics of the genus Bowenia show some similarity with those of the latter group. Except for two species of Macrozamia which are narrowly elliptic, all of the genera have widely elliptic pollen and share a psilate exine surface and the thinnest sexine with nearly identical arrangement of alveoli. Pollen grains of the species in the genus Dioon exhibit a unique morphology but are more similar to Stangeria than they are to those of taxa in Zamiaceae. The circular outline of the grains and the foveolate exine surface are characters shared by these two genera, but several morphological features distinguish Dioon from Stangeria. Ceratozamia and Zamia share a widely elliptic shape, foveolate exine surface and nearly identical sexine, as well as morphological features and chromosome numbers. They differ from Microcycas in sexine thickness, gross morphology and chromosome numbers. The pollen grains of Cycas circinalis and C. revoluta differ in size and structure of the sexine from all other genera and from each other, substantiating their distinct subgeneric delimitations.     

THE POLLEN WALL OF CANNA AND ITS SIMILARITY TO THE GERMINAL APERTURES OF OTHER POLLEN

The pollen wall of Canna generalis Bailey is exceptionally thick, but only a minor part of it contains detectable amounts of sporopollenin. The sporopollenin is in isolated spinules at the exine surface and in the intine near the plasma membrane. There is no sporopollenin in the > 10 μ thick channeled region between spinules and intine. We suggest that the entire pollen wall of C. generalis is similar to the thick intine and thin exine typical for germinal apertures in many pollen grain types. Considered functionally, the Canna pollen wall may offer an infinite number of sites for pollen tube initiation and would differ significantly from grains that are inaperturate in the sense of an exine lacking definite germinal apertures.     

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 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 EVOLUTION OF POLLEN TETRADS IN ONAGRACEAE

In Onagraceae, pollen is shed in mature tetrads in most Epilobieae, many species of Ludwigia (Jussiaeeae), and two closely related species of the large genus Camissonia (Onagreae). Mature tetrads of Camissonia cardiophylla and representative species of Epilobium and Ludwigia were examined with light, scanning, and transmission electron microscopes. Morphological diagnoses of monad units indicated that individual taxa could be readily distinguished. Statistical analyses of tetrads which remained after acetolysis treatment revealed significant differences in the strength of the binding mechanisms. Mechanisms of tetrad cohesion were found to consist of two principal types. Common to all taxa is cohesion of pollen wall surfaces at the aperture margins; this mechanism is well known in many angiosperm groups. With the exception of Camissonia, the remaining taxa also display binding by means of short exine fragments between adjacent pollen units. These fragments, termed bridges and reported here for the first time, are located in the area extending from the aperture margins to near the center of the proximal exine faces. Thin sections reveal that layers of the bridges are identical with those of the exine. Comparisons were made between bridges and viscin threads, both of which occur on the proximal faces of the grains. Viscin threads are present on all pollen grains in Onagraceae and exhibit distinctive morphologies, and bridges were viewed morphogenetically as related to viscin threads but including an endexine layer and occupying a position near the apertures where cohesion of wall surfaces also occurs. In an evolutionary sense, the formation of mature tetrads almost certainly occurred independently in Camissonia and may have done so in Ludwigia and the Epilobieae.     

POLLEN APERTURE POLYMORPHISM AND GAMETOPHYTE PERFORMANCE IN VIOLA DIVERSIFOLIA

Pollen aperture polymorphism is studied in Viola diversifolia, where all plants produce three- and four-apertured pollen grains. We tested whether there are genetic differences among plants for the proportions of the different pollen morphs, and whether the morphs differ in gametophytic performance. Results show that the more apertures a pollen grain has, the more quickly it germinates but that few-apertured pollen grains have faster growing pollen tubes and longer life expectancies. The proportions of the different pollen morphs, together with pollen tube growth rates, may be inherited traits based on differences among maternal families. These results suggest that the different pollen morphs are favored in different pollination ecology situations. The production of several pollen morphs by the same individual could therefore be evolutionarily advantageous.     

POLLEN TETRADS IN STENOMESSON ELWESII (AMARYLLIDACEAE)

Pollen tetrads within Amaryllidaceae are reported for the first time for Stenomesson elwesii. Tetrads were examined with light and scanning and transmission electron microscopy. The tetrads are tetragonal in shape. Tetrad-members are rounded-triangular in equatorial view and elliptic in polar view. Average polar diam is 50.1 μm; average longest equatorial diam is 62.5 μm. Morphology of the aperture is monosulcate. Exine sculpturing of the intectate pollen is gemmate. Systematic implications are briefly discussed.     

A NEW STRUCTURALLY PRESERVED PENNSYLVANIAN CORDAITEAN POLLEN ORGAN

Permineralized specimens of the pollen organ Gothania (Hirmer) consist of a primary axis bearing pollen cones in the axils of bracts that are four ranked. The bilaterally symmetrical primary axis consists of a uniform parenchymatous pith surrounded by up to 15 endarch-mesarch axile bundles. The cortex is two-parted and consists of an inner zone of subepidermal fibers. Bract traces arise from the ends of the ellipsoid stele. Traces to the cones are derived from the open ends of the stele, and at higher levels form a centrarch-medullated vascular system. Each pollen cone is constructed of up to 25 helically arranged scales, each vascularized by a single trace that may dichotomize. Scales are elongate and broad, and histologically composed of mesophyll parenchyma and fibrous layers. Stomata are restricted to the adaxial surface between rows of fibers. Up to 10 distal scales may be fertile, each with 4 elongate pollen sacs at the tip. Large monosaccate grains of the Felixipollenites-type are densely packed in each pollen sac. The well-preserved specimens of Gothania provide an opportunity to compare this genus with pollen cones assigned to the genus Cordaianthus, and to relate isolated plant organs to the Cordaitales.     

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.     

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

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

POLLEN MORPHOLOGY 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.     

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.     

FINE STRUCTURE AND CYTOCHEMISTRY OF LILIUM POLLEN TUBES

Growth of the Lilium longiflorum pollen tube in vitro is restricted to a zone extending back 3–5 μ from the tip. Electron micrographs of cross and longitudinal thin sections of L. longiflorum and L. regale pollen tubes reveal that the cytoplasm of the nongrowing region of the tube contains an abundance of mitochondria, amyloplasts, Golgi bodies, endoplasmic reticulum, lipid bodies, and vesicles. In contrast, the growing tip is characterized by an abundance of vesicles and an absence of other cytoplasmic elements. The vesicles appear to be of 2 types. One is spherical, about 0.1 μ in diameter, stains strongly with phosphotungstic acid, apparently arises from the Golgi apparatus and appears to contribute to tube wall and plasmalemma formation. The other type is irregular in shape, 0.01-0.05 μ in diameter, stains strongly with lead hydroxide, and is of unknown origin and function. Cytochemical analysis indicates that the tips of L. longiflorum pollen tubes are singularly rich in ribonucleic acid, protein, and carbohydrate. These findings are discussed in relation to tube growth.     

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.     

ULTRASTRUCTURE AND DEVELOPMENT OF MESOZOIC POLLEN: CLASSOPOLLIS

Because of its complexity one of the most unusual fossil pollen types is the genus Classopollis. Grains of this broadly defined Mesozoic taxon range from the late Triassic into the Cretaceous (Turonian), and include forms that are spherical with a subequatorial rimula. On the proximal pole is a trilete mark, and on the distal surface a thin area in the sporoderm termed the cryptopore. Ultrastructural studies of Classopollis have been completed on grains extracted from the pollen cone Classostrobus comptonensis collected from the Lower Cretaceous Wealden beds on the Isle of Wight, England. The sporoderm consists of clearly defined nexine and sexine components, with the mature nexine composed of approximately 20 electron dense lamellae, each about 10 nm thick. The sexine consists of four (S_1–4) easily recognizable layers, with the most prominent zone formed of coarse, inwardly-tapering elements. The S₂ layer is uniformly thickened, except in specialized areas (e.g., trilete, rimula, cryptopore) where it becomes thin. The remaining wall layers include spinules that ornament the surface and a uniform series of small lacunae associated with the spinule bases. The presence of orbicules and a complex system of membranes associated with the grains extracted from less mature cones provides an opportunity to trace some developmental stages in Classopollis sporoderm ontogeny, and to compare these stages with those of selected extant pollen types. The functional significance of the infrastructure in Classopollis pollen is discussed.     

POLLEN SUCCESSION,ABSOLUTE POLLEN FREQUENCY,AND RECURRENCE SURFACES IN CENTRAL JAPAN

A generalized pollen diagram, associated with three simplified pollen diagrams from Tsubogakure (1,500 m), Nakatashiro (1,400 m) and Yashima-ga-hara (1,650 m) bogs in central Japan, represents five pollen zones for the last 12,000 years, i.e., L (before 10,500 B.P., pine-boreal); R I (10,500–9,500 B.P., boreal-oak); R II (9,500–4,500 B.P., temperate); R IIIa (4,500–1,500 B.P., temperate-boreal); and R IIIb (after 1,500 B.P., pino-nonarboreal). Absolute pollen frequency is high in well-humified peat, in gyttja (deposited immediately below the first peat formation), and in peat in the part of the R IIIa cooling stage, and it is low in less-humified peat, since the former sediments have slow and the latter rapid rates of accumulation, respectively. Changes of plant communities should affect the absolute pollen frequency, but changes of sedimentation rates are more critical. After a re-examination of Nakatashiro bog, it is hardly possible to recognize a recurrence surface in the European sense, nor can the accumulation of thin tephra on the bog surface be responsible for a temporary halt in peat growth. Nakatashiro bog (like Tsubogakure bog) is a low-moor bog, and true recurrence surfaces would not be expected in it. In a raised Yashima-ga-hara bog, however, there are seven recurrence surfaces without a volcanic ash band; the well-humified horizons correspond to low values of Sphagnum spores and Cyperaceae pollen, and the less-humified horizons to high values of Sphagnum spores and Cyperaceae pollen. Peaks of these species in the upper core correspond to those Cryptomeria pollen, an indicator of the high precipitation (generally >1,800 mm/yr). Regularly interrelated fluctuations among these species, absolute pollen frequency, and peat horizons suggest that the precipitation was low in well-humified peat and was high in less-humified peat.     

POLLEN ULTRASTRUCTURE OF THE TRIBE INGEAE (MIMOSOIDEAE: LEGUMINOSAE)

All genera within the Ingeae, excluding Wallaceodendron, were examined with the transmission electron microscope. Thin sections reveal two pollen types (Types I and II) distinguished primarily by differences in polyad cohesion and ektexine organization. Type I polyads (only eight-grained species of Calliandra) are calymmate and the ektexine of individual cells is continuous around the grain, organized into a thin, foraminate tectum, irregularly shaped, often basally flared, foraminate columellae and thin, discontinuous foot layer. Type II polyads (16-grained species of Calliandra and remaining Ingeae) are predominantly acalymmate with individual grains typically free from one another or rarely, partially calymmate, i.e., individual grains show limited forms of attachment through small endexinous bridges (Pithecellobium latifolium [Zygia], Lysiloma) or localized appression of adjacent endexines (Pithecellobium daulense [Cathormion]). The adhesion of individual grains through localized fusion of lateral-distal and proximal ektexine in Enterolobium is unique among the partially calymmate Type II polyads. Ektexine in Type II polyads, largely restricted to the distal face, is composed of a thick, channeled tectum, granular interstitium and when present, thin discontinuous foot layer. Lateral-distal and proximal areas exhibit only endexine and, occasionally, a foot layer. The occurrence of nondistal ektexine is restricted to Enterolobium. The pollen data suggest that the acalymmate Ingeae polyads composed of grains with porate apertures, thick, highly channeled tectum, granular interstitium and lack of, or greatly reduced foot layer, are clearly derived within the Mimosoideae. Type I calymmate polyads appear to be independently derived. Ultrastructural data suggest that the Ingeae, excluding the eight-grained Calliandra species, represent a natural grouping with a close affinity to the Acacieae.