PALYNOLOGY AND SYSTEMATICS OF CUPHEA (LYTHRACEAE). II. POLLEN MORPHOLOGY AND INFRAGENERIC CLASSIFICATION

Present taxonomic studies in Cuphea (Lythraceae) reveal that a broad spectrum of biosystematic data is required for an understanding of evolutionary relationships among this large and complex group of species. As part of these studies pollen data have proven to be of greater pragmatic value than is frequently true for more stenopalynous taxa. A survey is made of pollen types in 160 species of Cuphea with special reference to specific taxonomic problems, and a summary presented of points at which pollen data can profitably contribute to taxonomic revision of the genus.     

PALYNOLOGY AND SYSTEMATICS OF THE LYTHRACEAE. I. INTRODUCTION AND GENERA ADENARIA THROUGH GINORIA

A palynological survey, including LM, SEM, and TEM, is presented for 10 genera of the Lythraceae s.s. (plus Alzatea to be assigned separate family status): Adenaria, Ammannia, Capuronia, Crenea, Cuphea, Decodon, Didiplis, Diplusodon, Galpinia, and Ginoria. Discussion of the results is deferred until the survey is complete, but already information of taxonomic interest is available for the 11 genera studied. The pollen data, consistent with results from cytology, anatomy, and embryology, suggest the current subgeneric classification into the Tribes Lythreae and Nesaeeae based on complete or incomplete septation of the ovary is not valid. The pollen of Alzatea is a generalized type and lacks the morphological features most characteristic of the Lythraceae (pseudocolpi, sculptured exine). The genus is being described as a separate family, and the pollen morphology is consistent with this treatment. The pollen of Cuphea is more diverse and shows little similarity to other genera in the family. It also has a unique combination of floral characters and lipid chemistry that suggests a separate category within the developing subfamilial classification of the Lythraceae.     

Pollen morphology in the genus Coffea (Rubiaceae) and its taxonomic significance

The pollen morphology of 38 representative species and varieties within the genus Coffea L. is described. Eight pollen types, placed in two major groups, have been identified on the basis of the number of colpi, colpus characteristics, exine morphology and pollen size. Differences in pollen structures of the species studied do not correspond to present taxonomic groupings, but support a current proposal to place some of the species in the genus Paracoffea Leroy. Among the four sections in the genus, the section Eucoffea contains a high degree of pollen polymorphism.     

POLLEN MORPHOLOGY AND ULTRASTRUCTURE OF KRAMERIA (KRAMERIACEAE): UTILITY IN QUESTIONS OF INTRAFAMILIAL AND INTERFAMILIAL CLASSIFICATION

The genus Krameria is currently recognized as an enigmatic, monotypic family of dicotyledons. Previous studies of morphology, anatomy, and cytology have been unable to establish unequivocably its phyletic affinities. We report here the results of an intensive investigation of the pollen of Krameria using light, scanning electron (SEM), and transmission electron microscopy (TEM). Pollen characteristics of the genus were compared to those of all families referred to the Polygalales and to selected species of the Leguminosae-Caesalpinoideae, both groups with which Krameria has historically been allied. Superficially, the pollen of Krameria resembles that of the legumes more than that of genera in the Polygalales. However, in ultrastructure, it differs from the pollen of all taxa investigated from both groups. Within Krameria, several variations of a basic type of 3-colporate pollen are discernible. Species with similar pollen variants appear, on the basis of other morphological data, to represent natural groups within the genus. Nevertheless, an arrangement of groups of species of Krameria from “least” to “most” specialized, based on a logical sequence of modification of the pollen morphology, does not agree with any sequence of specialization using other morphological or ecological characters. It is concluded that pollen morphology and ultrastructure has systematic value for intrafamilial groupings of the Krameriaceae but that palynological modifications are incongruous with trends of specializations of other characters and, like many other lines of investigation, pollen studies do not provide significant data as to the phylogenetic affinities of the family.     

Pollen morphology of Chamaebuxus (DC.) Schb., Chodatia Paiva and Rhinotropis (Blake) Paiva (Polygala L., Polygalaceae)

Polygala L. is a large and highly diverse genus with complex taxonomy, but pollen morphological information for this taxon is scarce. In the present study, pollen characters have been used to assess the taxonomic delimitation and phylogenetic relationships of three newly established subgenera of Polygala: Chamaebuxus, Chodatia and Rhinotropis (sensu Paiva). The pollen morphology of 22 species has been examined using light microscopy and scanning electron microscopy of acetolysed material. The pollen of 15 of the species is examined for the first time. The pollen grains are isopolar, radially symmetrical, tectate and, typically, polyzonocolporate with numerous colpi running parallel to the polar axis, and an endocingulum around the equator. Two pollen types can be distinguished: Type I, which includes species belonging to Rhinotropis, and Type II, which includes species from Chamaebuxus and Chodatia. The two pollen types are described and the pollen of the three studied subgenera is illustrated. Despite the low infrageneric morphological diversity observed within the genus Polygala, quantitative characters of pollen grains support the current classification of the subgenera Chamaebuxus, Chodatia and Rhinotropis, and reveal a closer relationship between the first two taxa. Pollen characters are shown to be a useful and informative tool for assessing taxonomic position and phylogenetic relationships within Polygalaceae, especially at higher taxonomic levels.     

A SYNOPSIS OF NORTH AMERICAN ASTERS: THE SUBGENERA,SECTIONS AND SUBSECTIONS OF ASTER AND LASALLEA

Traditional treatments of Aster were found unsatisfactory in predicting cytological and morphological characteristics of members of various subgenera, sections and subsections of different authors. A realignment of species into these infrageneric categories is presented and is based on sets of shared morphological characteristics of roots or rhizomes, leaves, and phyllaries. Eight subgenera are recognised in Aster. Subgenus Aster is divided into five sections on morphological characteristics: three have X = 9, one has X = 8 and one has X = 7. Subgenus Aster section Dumosi has been redefined and contains all the X = 8 species. Ten X = 5 species and one X = 4 species previously classified in up to six sections of subgenus Aster were found to be morphologically and cytologically related to each other, but unrelated to the other species of Aster. These eleven species have been assigned to the genus Lasallea sensu Semple & Brouillet. Lasallea has been divided in sections and subsections with new combinations presented in this paper.     

PALYNOLOGY AND SYSTEMATICS OF THE LYTHRACEAE. II. GENERA HAITIA THROUGH PEPLIS

Pollen morphology for 12 genera of the Lythraceae is described, using light microscopy (LM), scanning (SEM) and transmission electron microscopy (TEM). The genera surveyed are Haitia, Heimia, Hionanthera, Lafoensia, Lagerstroemia, Lawsonia, Lythrum, Nesaea, Orias (= Lagerstroemia), Pehria, Pemphis, and Peplis. Among results of taxonomic significance are 1) the similarity of pollen between Haitia and Ginoria, suggesting that future classifications reflect a close relationship between these genera, 2) a comparable similarity in pollen between Pehria and Adenaria, 3) the distinction of Hionanthera pollen from that of Ammannia, 4) an exine similar in certain ultrastructural characteristics between Orias and some species of Lagerstroemia, although in general aspect they appear quite different, 5) that within the Didiplis-Lythrum-Peplis complex, there are three palynologically distinct groups, with Didiplis standing apart from the somewhat more similar, but nevertheless, distinct Lythrum and Peplis, and 6) that considerable differences exist between pollen of Pemphis acidula and Pemphis madagascariensis, supporting removal of the latter species to a new genus. This pollen survey is part of a multi-disciplinary effort to clarify generic limits in the Lythraceae, and evaluate relationships within the family and among families of the Myrtales.     

CHROMOSOME NUMBERS IN CUPHEA (LYTHRACEAE): NEW COUNTS AND A SUMMARY

The American genus Cuphea with ca. 260 species is extremely diverse with respect to chromosome number. Counts are now available for 78 species and/or varieties, or 29% of the genus. Included in this study are first reports for 15 taxa from Brazil, Cuba, Dominican Republic, Mexico, and Venezuela. Twenty-two different numbers are known for the genus, ranging from n = 6 to n = 54. The most common number in the primary center of species diversity in Brazil is n = 8, which is regarded as the base number of the genus. Two numbers are most common in the secondary center in Mexico, n = 10 and n = 12. Species with n = 14 or higher are considered to be of polyploid origin. Polyploids comprise 46% of the total species counted and appear in 9 of the 11 sections for which chromosome numbers have been reported. Aneuploid species comprise ca. 25% of the genus and are known from 7 of the 11 sections. The two subgenera are not characterized by different chromosome numbers or sequences of numbers. None of the 14 sections are circumscribed by a single chromosome number. Morphological and ecological variability in widespread, weedy species is correlated with differing chromosome numbers in some species whereas in others the chromosome number is stable. Summary of chromosome numbers by taxonomic section is presented. Section Euandra, centered in eastern Brazil, and the largest section of the genus, appears to be chromosomally most diverse. In section Trispermum, characterized by difficult, variable species with intermediate forms, two of the four species studied have polyploid races. Section Heterodon, endemic to Mexico and Central America and comprising most of the annual species of the genus, is best known chromosomally. Chromosome numbers have been counted for 25 of 28 species, and 12 different numbers are reported. The most advanced sections, Melvilla and Diploptychia, with numerous species occurring at higher altitudes, are characterized by high polyploids. Apomictic species occur in sect. Diploptycia. The cytoevolution of Cuphea is complex with frequent polyploid and aneuploid events apparently playing a significant role in speciation in both centers of diversity.     

Pollen morphology of Astragalus section Hymenostegis (Fabaceae) and evaluation of its systematic implications

Astragalus is with nearly 3000 described species the largest genus of flowering plants. So far analyses of pollen characters have only been conducted for a few species of the groups within the genus. Here we analyse pollen grains of 22 species representative for Astragalus section Hymenostegis using scanning electron microscopy. We found the basic shape of the pollen grains to be oblate-spheroidal and apertures to be tricolpate as for other eudicots. The sculpturing pattern of the exine is micro-reticulate. Pollen grains show low morphological variation among different species of this section, but differences occur between sections of the genus. We conclude that the vast morphological differentiation that occurred during the rapid radiation of section Hymenostegis was not accompanied by comparable differentiation in pollen morphology.     

Pollen morphology within the Monodora clade,a diverse group of five African Annonaceae genera

Pollen morphology has played a major role in elucidating infrafamiliar‐level systematics and evolution within Annonaceae, especially within the African genera. The Monodora clade is composed of five genera, Asteranthe, Hexalobus, Isolona, Monodora and Uvariastrum, which are restricted to Africa and contain together c. 50 species. A molecular phylogeny of the family showed that the monophyly of the Monodora clade is strongly supported and that it is part of a larger clade of 11 African genera. In order to support classification a detailed survey was made of the pollen morphological variation within the Monodora clade, using scanning and transmission electron microsopy. For the two most species‐rich genera, Isolona and Monodora, a molecular species‐level phylogeny was used to assess the taxonomic usefulness of the pollen characters. The survey showed a wide range of pollen morphological diversity. The most conspicuous variation concerned the occurrence of monads without a thicker outer foliation in the basal exine layer in Isolona in contrast to tetrads with a thicker outer foliation in Asteranthe, Hexalobus, Monodora and Uvariastrum. At the infrageneric level, Hexalobus, Isolona and Monodora showed the largest diversity, with various pollen types based on tectum morphology. Hexalobus is exceptional with three types within only five species. The pollen types defined in this study are hardly useful in characterizing major groups identified within both Isolona and Monodora, but they do illustrate relationships within smaller groups.     

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 its phylogenetic implications in the genus Picea

Pollen grains from 20 spruce species were examined using scanning electron microscopy, to study the phylogeny of pollen morphology in spruce. Morphological characteristics of the pollen were examined among different species based on statistical data and photographic observation. The mean pollen size of 20 spruce species ranged from 84.327 to 118.793.18 μm. Based on morphological photography, two zygomorphic bladders were distributed on the underside of the corpus and were found at the edge of the cap near the corpus surface. The bladder’s surface texture was smooth and had no distinct variations among species. Germinal furrows formed clear striates between the two bladders. According to clustering analysis, 20 spruce species were classified into three sections, namely, Castict, Schrenkiana and Picea, and the Picea section was classified into two subsections, Jezonensis and Picea. Corpus veins experienced notable differentiation among sections, and their evolutionary trend progressed through sections Castict, Schrenkiana and Picea. Within subsection Picea, parallel evolution occurred between the two subsections.     

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.     

TYPES OF LATICIFERS AND CRYSTALS IN JATROPHA AND THEIR TAXONOMIC IMPLICATIONS

Anatomical examination of 37 species of Jatropha (Euphorbiaceae) has revealed the occurrence of two distinct types of laticifers—articulated, and nonarticulated—in addition to idioblasts that are irregularly shaped individual cells. With the notable exception of J. augustii Pax & K. Hoffm. (section Peltatae (Pax) Dehgan & Webster), these idioblasts occur in the two most advanced sections of the subgenus Curcas (Adans.) Pax, namely Loureira (Cav.) Muell. Arg. ex Pax and Mozinna (Ortega) Pax. The presence of two laticifer types and “idioblastic laticifers” in the genus, in association with the morphological reduction series found with evolutionary advancement are, therefore, significant in the delimitation of sections and subsections. Further evidence is presented by the occurrence of “chambered crystalliferous cells” in subgenus Curcas, but not in subgenus Jatropha (=Adenorhopium Griseb.). Reexamination of laticifers in other genera of the Euphorbiaceae is suggested as a possible means of alleviating the long-standing taxonomic dilemma of this large and morphologically diverse family.     

Studies in Montia L. and Claytonia L. and Allied Genera

The pollen morphology of several genera in Portulacaceae is described. Particular attention has been paid to the genera of the subfamily Montioideae, as a stage of continued monographical studies. Among genera especially dealt with are Claytonia, Montia, Crunocallis, Naiocrene, Neopaxia, Mona, Maxia, Limnalsine, and Montiastrum. In the taxonomical treatment of these genera the pollen morphology has proved to afford many important additional characters.

The pollen grains of Claytonia are distinguished from those of the remainder in being 3-colpate. The grains of the Claytonia-type have many similarities with those of Lewisia, a genus of the subfamily Portulacoideae. The other genera of Montioideae have pantocolpate pollen grains. Among these genera several different pollen types are distinguished, chiefly with regard to the sexine structures and the aperture membranes. The Montiastrum-type is especially interesting, with tholate grains, a particular pollen type not met with in any other genus in the family. The pollen morphology of some genera in the Portulacoideae is also treated. In some species in Calandrina and Talinum pantotreme pollen grains are observed with apertures transitional between pori and colpi. The apertures of the pantotreme grains are arranged in characteristic patterns.

Particular attention has been given to the variation of the pollen morphological characters. This variation has been examined with regard to the differences between different populations of the same species as well as between different species. The greatest variation has been observed in the shape and size of the grains. The structure and sculpture and thickness of the sexine and the aperture membranes are less variable. Some polyploid taxa are connected with the occurrence of pollen grains with divergent and varying aperture numbers.

In a survey of the genera the taxonomical results of the investigation are presented with particular regard to the pollen morphology. The new genus, Maxia Ö. Nilss., is described. One new species, Montia clara Ö. Nilss., is described and some new combinations are made.

Pollen morphological diagnoses are given for 46 different taxa. The aperture conditions for 96 different species are presented.     

秦巴山区悬钩子属植物的地理分布及花粉形态观察

秦巴山区有悬钩子属植物40种15变种,隶属于3组15亚组,是我国悬钩子属的一个集中分布区,在该属植物起源和演化中占重要地位。部分类群的花粉形态观察结果表明孢粉学分析是该属植物系统分类研究的一个有效手段。     

中国赤车属新分类

(1)根据该文中赤车属与楼梯草属各种形态学的区别,认为赤车属较为原始,在楼梯草属之前自冷水花属衍生而出。(2)将赤车属划分为5组;将C.B.Robinson描述的Elatostemoides属降为组级群,作为赤车属中接近原始群全缘赤车组sect.Pileoides的一个组处理,此外还描述了一个与sect.Elatostemoides近缘的1新组,羽脉赤车组sect.Leiolaena,并根据叶、叶的脉序、雌花序、雌花被片和瘦果等器官的演化趋势给出显示赤车属属下诸群间亲缘关系的系统发育树。(3)对中国赤车属的属下分类群进行了修订,共承认32种1亚种7变种,给出检索表、描述及多幅插图,并将它们划分为4组、2亚组和9系,其中除sect.Leiolaena之外,还新描述了2亚组、5系、2种和3变种;在2003年被归并为同物异名的长柄赤车、曲毛赤车、波缘赤车、小赤车和富宁赤车的种级地位得到恢复。此外,还写出了中国赤车属的分类学简史和地理分布。