The endotegmen tuberculae: an account of little-known structures from the seed coat of the Hydrocharitoideae (Hydrocharitaceae) and Najas (Najadaceae)

SHAFFER-FEHRE, M., 1991. The endotegmen tuberculae: an account of little-known structures from the seed coat of the Hydrocharitoideae (Hydrocharitaceae) and of Najas (Najadaceae). The sub-cellular protuberances from the inner tegmen layer have been given various names in earlier literature. Here named 'endotegmen tuberculae', they vary in shape, size and density of distribution in the endotegmen of different taxa. These characteristics make them useful for taxonomic research at both genus and species level. Their shape is a major consideration in transferring Hydrocharis dubia to the genus Limnobium. A further structure, 'exotegmen tuberculae', has been observed for the first time. This paper gives a survey of these structures and illustrates their taxonomic importance and its implications.     

中国茨藻科植物种皮微形态特征及其系统学意义

本文应用扫描电镜对国产茨藻科植物的外种皮和内种皮微形态特征进行了研究,结果表明茨藻科植物在种子表面细胞形态、表面纹饰和内种皮内层细胞形态及小瘤状突起的大小和密度等方面存在差异,可作为种级水平分类的依据,并首次发现外种皮微形态特征也与内种皮内层小瘤状突起一样具有重要的系统学意义。根据种皮微形态特征的研究结果,我们支持Miki(1937)和Shaffer-Fehre(1991b)等关于茨藻科与水鳖科近缘,而不应放在眼子菜目的观点。对弯果茨藻Najas ancistrocarpa A．Br．ex Magnus和草茨藻N.graminea Del.外种皮的研究结果表明其外种皮细胞表面具瘤状颗粒而非网状加厚。     

Phylogeny of the family hydrocharitaceae inferred fromrbcL andmatK gene sequence data

The family Hydrocharitaceae, with 15 genera and ca. 80 species, shows a remarkable morphological diversity which presumably developed as an adaptation to their aquatic habitat. This is particularly true in the case of the many different kinds of pollination mechanisms. To gather more basic information regarding the adaptive evolution of Hydrocharitaceae, we have carried out a phylogenetic analysis based on the sequences of therbcL andmatK. Our resulting neighbor-joining distance tree provides the following insights: (1) none of the previous classification systems were supported by molecular phylogenetic tree; (2) Najas (Najadaceae), which has never been included in Hydrocharitaceae except in Shaffer-Fehre's (1991) system based on seed coat structures, is an ingroup of Hydrocharitaceae; (3) Limnocharitaceae and Alismataceae are sister groups of Hydrocharitaceae; (4) the three marine genera,Halophila, Enhalus andThalassia, are monophyletic; and (5) a peculiar pollination mechanism specific to Hydrocharitaceae (Hydrocharitaceae-epihydrophily), underwent a parallel evolution.     

中国水鳖科植物种皮微形态特征及其系统学意义

通过扫描电镜对国产水鳖科植物(包括6属13种)的种皮微形态特征进行观察,并作了系统描述。根据种皮细胞形态、外种皮表面纹饰和内种皮内层小瘤状突起的特点将水鳖科植物的种皮微形态特征划分为3种类型,即海菜花型(海菜花属)、水鳖型(水鳖属)和苦草型(苦草属、水筛属、虾子草属和黑藻属),并作出了分属检索表。本文结果表明,种皮微形态特征可作为该科族、属以及属内种级水平分类的依据,对探讨属间关系和该科的系统发育关系亦具有重要的价值。种皮微形态特征支持Hutchinson(1959)和Eckhardt(1964)将海菜花属和水鳖属分别作为一个独立的族处理的观点。苦草属、水筛属和虾子草属种皮微形态特征的高度相似性表明它们间有密切的联系,不支持将它们置于不同亚科和族的分类处理。黑藻属虽与上述3属近缘,但其外种皮特征则较为独特,因此与水筛属放在不同族中更为合理。本文种皮微形态特征的研究结果支持iki1937)和Shaffer-Fehre(1991b)等关于水鳖科与茨藻科近缘的观点。     

The problematic systematic position of Ceratostratiotes Gregor (Hydrocharitaceae ?) – morphological, anatomical and biochemical comparison with Stratiotes L.

 The enigmatic fossil taxon Ceratostratiotes sinjanus (Kerner) Bužek has been considered as either a Hydrocharitaceae seed (monocot) or a Ceratophyllaceae fruit (dicot). The co-occurrence of seeds of Stratiotes kaltennordheimensis (Zenker) Keilhack (Hydrocharitaceae) and Ceratostratiotes in Early Miocene sediments in Langau (Lower Austria) enabled comparisons of morphological-anatomical features and lignin compositions, independent of diagenetic biases. The biochemistry of the Ceratostratiotes seed coat wall is not monocotyledon-like and is dissimilar to that of the co-occurring Stratiotes testae. In contrast, the Ceratostratiotes seed coat anatomy resembles that of Hydrocharitaceae genera with the micropyle and raphe being very similar to those in seeds of Stratiotes. Although the horizontal arrangement of the Ceratostratiotes spines is different from the longitudinal arrangement of the surface ridges in Stratiotes kaltennordheimensis and the spines in Blyxa (Hydrocharitaceae), the spine structures are very similar. A mixture of similarities and differences characterizes the morphological-anatomical features. Thus the systematic affinity of Ceratostratiotes still remains enigmatic. Received April 18, 2002; accepted October 18, 2002 Published online: February 4, 2003     

Seed coat micromorphology of South American species of Hybanthus (Violaceae)

The seed‐coat of several species of Hybanthus subg. Ionidium from South America were studied by SEM (scanning electronic microscopy). Three different levels of sculpture were observed on the seed coat. The micro‐morphological patterns showed great variation between the species and should be considered as additional characters in future taxonomic treatments. The presence of a conspicuous elaiosome was observed; this structure is well known in the genus Viola and is most likely related to seed dispersal mediated by ants.     

The development of the fruits and seeds of campomanesia (Myrtaceae)

The glandular nature of the “seed coat” has long been the most important taxonomic character defining the genusCampomanesia. It is shown here that the structure, which has been taken for the “seed coat,” is actually the locular wall. The development of the ovaries and seeds ofCampomanesia is discussed and illustrated. Three characteristics ofCampomanesia are identified as distinguishing it from other genera of the Myrtaceae: (1) the abortion of all but one ovule in each fertile locule; (2) the protective, glandular locular wall that serves as a false seed coat; and (3) the large number of locules in the ovaries. Reasoning is presented to explain the possible sequence of evolutionary origin of these three characteristics.     

SYSTEMATIC SIGNIFICANCE OF SEED-SURFACE FEATURES IN ORTHOCARPUS (SCROPHULARIACEAE—SUBTRIBE CASTILLEJINAE)

Scanning electron microscope and light microscope examination of seed-coat features of 26 species of Orthocarpus have allowed recognition of many species-level differences (summarized in a key) and of three seed-coat types that parallel taxonomic subgroups but support realignments at generic and infrageneric levels. Type 1 seeds (subg. Orthocarpus, sect. Orthocarpus) have a lateral hilum, sculptured inner tangential seed-coat walls, and a tightly fitting outer seed coat. They are very similar to seeds of Cordylanthus. Seeds of Types 2 and 3 have a terminal hilum and membranous inner tangential cell walls. Type 2 seeds (subg. Orthocarpus, sects. Castillejoides and Cordylanthoides, with one exception) have a net-like, loosely fitting outer seed coat that shows close relationship to seeds of Castilleja. Inner tangential walls of Type 2 seeds normally rupture. Type 3 seeds (subg. Triphysaria, with two exceptions) have a tightly fitting outer seed coat and inner tangential walls are always retained. Seed features support evidence from floral morphology and chromosome numbers that Orthocarpus as currently recognized is not a monophyletic lineage.     

Seed and trichome morphology of the Egyptian Fagonia (Zygophyllaceae) with emphasis on their systematic implications

Seed and trichome morphology of 13 taxa representing the genus Fagonia in Egypt have been studied using light (LM) and scanning electron microscopy (SEM). Macro‐ and micro‐morphological characters, including seed shape, seed size, seed coat sculpture, trichome density and structure were studied to evaluate their systematic significance. Seed shape and trichome density were found to have only minor taxonomic value, but seed coat sculpturing and trichome structure proved to be valuable characters. These characters offer evidence to combine certain species, e.g. Fagonia kassasii and F. schimperi in F. bruguieri, Fagonia microphylla in F. scabra and Fagonia thebaica in F. arabica. However, F. sinaica, F. tristis, F. boveana and F. isotricha should be maintained as separate species. Generally, the different patterns of seed and trichome morphology are useful in delimitation of species within the genus Fagonia, but they could not be used to characterize groups of related species.     

Systematic relevance of seed coat anatomy in the European heathers (Ericeae, Ericaceae)

The anatomy of the seed coat of the European species of tribe Ericeae (Calluna, Daboecia and Erica) of the Ericaceae family was studied, and the taxonomic importance of their characters was analyzed. The seed coat is mostly formed by a one-cell layer with thick, pitted inner walls and thin outer walls that collapse at maturity over the inner walls. The cell junctions are either raised with anticlinal walls up to four times the height of the periclinal walls or are not raised with similar values for the height of both the anticlinal and periclinal walls. Three main cell junction types were found and described. The thickness of the inner walls is variable, but there is a large overlap among the results for different species. Calluna vulgaris is the only species with no pits, and E. multiflora has a pitted pattern on its inner walls, which is distinctive from the rest of the species. Our main results agree with the external seed morphology, and valuable new data were obtained for certain groups such as the E. cinerea-E. terminalis or the E. scoparia complex. The similarities that are found in seed coat characters are not in accordance with the classical taxonomic delimitation of infrageneric groups within Erica.     

Seed coat development in Leucospermum cordifolium (Knight) Fourcade (Proteaceae) and a clarification of the seed covering structures in Proteaceae

MANNING, J. C. & BRITS, G. J., 1993. Seed coat development in Leucospermum cordifolium (Knight) Fourcade (Proteaceae) and a clarification of the seed covering structures in Proteaceae . The development of the seed coat and pericarp is studied in Leucospermum cordifolium from ovule to mature seed. The ovule and seed are characterized by a tegmic pachychalaza. The pericarp is adnate to the integuments from anthesis and remains unthickened to maturity. The outer integument forms the seed coat and the seed is endotestal: the outer epidermis becomes tanniniferous and the inner epidermis develops into a crystalliferous palisade. The inner integument degenerates at an early stage. Examination of the literature reveals that the crystal palisade layer of the outer integument has been erroneously assumed to constitute an endocarp. This finding indicates that a re-interpretation of all published information on the seed coat in indehiscent Proteaceae is necessary before any speculations on the phylogenetic significance of the seed coat can be entertained.     

果皮和种皮微形态特征在杜鹃花属系统学研究中的应用

采用扫描电子显微镜对国产杜鹃花属Rhododendron6个亚属的代表种和近缘类群杜香属Ledum杜香L.palustre的果皮和种皮微形态特征进行观察,对杜鹃花属植物果皮微形态特征进行了系统描述,并通过比较现存杜鹃花属植物和种子化石,新发现一些居间的种子类型。结果表明,果皮和种皮微形态特征具有一定的系统学意义。叶状苞亚属subgen.Therorhodion的叶状苞杜鹃R.redowskianum的果实有短而疏的刺毛,无气孔,种子为无翅类,扁平,外围轮廓长椭圆形。杜鹃亚属subgen.Rhododendron植物果皮为百合花杜鹃型,具有鳞片,气孔器散生于指状突起之间,与叶表皮的微形态特征一致,其种子为百合花杜鹃型,表面具宽而浅的沟,呈脑纹状,有别于无鳞类杜鹃花。常绿杜鹃亚属subgen.Hymenanthes果皮为云锦杜鹃型,其角质层表面不规则,无表皮毛,偶见气孔;种子为云锦杜鹃型。映山红亚属subgen.Tsutsusi果皮为岭南杜鹃型,密生长表皮毛,角质层致密;种子为南边杜鹃型和岭南杜鹃型。微形态特征与“常绿杜鹃亚属和(国产)映山红亚属均为内部支持率很高的单系类群”的分子系统发育研究结果一致。马银花亚属subgen.Azaleastrum的马银花组sect.Azaleastrum和长蕊杜鹃组sect.Choniastrum微形态区别明显,支持各自为独立的单系类群。羊踯躅亚属subgen.Pentanthera的羊踯躅R.molle果皮特征明显,可与其他亚属区分,但种子类型更接近常绿杜鹃亚属。本研究结合分子系统发育资料和叶表皮微形态特征讨论了一些近缘类群的系统发育关系;研究结果支持将腺萼马银花处理为独立的种。     

The embryology and taxonomic relationships of Bretschneidera (Bretschneideraceae)

TOBE, H. & PENG, C.-I, 1990. The embryology and taxonomic relationships of Bretschneidera (Bretschneideraceae). We present the first report on the embryology of Bretschneidera , the only genus of Bretschneideraceae (which are one of 15 glucosinolate-producing families), to clarify its relationships. Embryologically Bretschneidera is characterized by the following features: ovule campylotropous, bitegmic and crassinucellate; outer integument thick, multiplicative and vascularized; embryo sac formation of the Allium type; seed exalbuminous; seed coat 'exotestal' with a palisade of columellar, thick-walled exotestal cells; mesotesta thick with the inner half aerenchymatous. These features suggest that Bretschneidera is distinct from any of the taxonomically related families but resembles both Hippocastanaceae and Sapindaceae (Sapindales) more closely than Moringaceae or Capparaceae (Capparales) which have been considered alternative allies, supporting most of the modern taxonomic treatments that place Bretschneidera as a separate family in Sapindales.     

A fossil Vallisneria plant (Hydrocharitaceae) from the Early Miocene freshwater deposits of the Most Basin (North Bohemia)

A new fossil species of Vallisneria (V. janecekii Bogner & Kva?ek, sp. n., Hydrocharitaceae) is reported from Early Miocene freshwater lake deposits of the North Bohemian brown coal basin (Czech Republic). It is based on a single fragmentary sterile plant bearing two clusters of ribbon-shaped widely serrulate leaves on a short rhizome. It is a potential producer of seeds described from equivalent and slightly older deposits in Saxony (Germany) as Vallisneria ovalis Mai.     

Isozymes as molecular markers for diploid and tetraploid individuals ofNajas marina (Najadaceae)

The genetic variability of five natural populations ofNajas marina L., i.e. one diploid of subsp.marina (Europe), two of subsp.intermedia (Europe) and both a diploid (C. Africa) and a tetraploid (Middle East) of subsp.armata, has been estimated by means of electrophoretic studies. These populations differ in their morphology and karyotype. Emphasis is placed on the characteristics and status of a tetraploid cytotype from Merkaz Sappir (Israel). Almost all the variation observed is expressed in seed alcohol dehydrogenase (ADH). The differences are in a unique allele of theAdh-2 locus and in the formation of novel heteromeric isozymes.Adh genes in seeds can be used as a marker for the autotetraploid character. The other enzyme systems tested failed in this respect. The genetic variability based on 23 loci is rather low. Nevertheless, the autotetraploid population has a higher or equal ratio of polymorphic loci than the related diploids. Cluster analysis illustrated not only thatNajas marina subsp.marina has diverged much from subsp.intermedia and subsp.armata, but also showed the difference between the latter two taxa, as well as the intermediate position of the autotetraploid population.     

Seed Coat Microsculpturing in Brassica and Allied Genera (Subtribes Brassicinae, Raphaninae, Moricandiinae)

To obtain new information on the taxonomy of Brassica and alliedgenera, seed surfaces of 44 species (78 accessions) belongingto 11 genera of subtribes Brassicinae, Raphaninae and Moricandiinaewere examined using a scanning electron microscope. Ten typesof basic ornamentation patterns were recognized. While fourtypes were represented by only one species each, six types hadmore than one species either representing one genus (two types)or more (four types). The different species in each of the sixtypes could generally be distinguished from each other on thebasis of differences in microsculpturing features. This analysisprovides evidence for the close relationship among the variousgenera within the subtribe Brassicinae, and also the closenessof Raphanus, Enarthrocarpus and Moricandia of subtribe Raphaninaeand Moricandiinae, respectively, with the Brassicinae. Copyright2000 Annals of Botany Company Brassicinae, Moricandiinae, Raphaninae, SEM, seed coat microsculpturing, taxonomic implications     

Ovules and seeds in Acalyphoideae (Euphorbiaceae): structure and systematic implications

Acalyphoideae, the largest subfamily of Euphorbiaceae, are investigated with respect to ovule and seed structure on the basis of 172 species of 80 genera in all 20 tribes of Acalyphoideae sensu Webster. All species of Acalyphoideae examined have bitegmic ovules with a non-vascularized inner integument. However, noticeable differences exist among and sometimes within the genera in the thickness of the inner and outer integument, the presence or absence of vascular bundles in the outer integument, whether ovules are pachychalazal or not, the presence or absence of an aril, seed coat structure (in terms of the best-developed mechanical cell-layer), and the shape of cells constituting the exotegmen. For the latter two characters, two different types of seed coat (i.e., "exotegmic" and "exotestal") and three different types of exotegmic cell (i.e., palisadal, tracheoidal and ribbon-like) were distinguished. Comparisons showed that three tribes Clutieae, Chaetocarpeae and Pereae are distinct from the other Acalyphoideae as well as from the other Euphorbiaceae in having an exotestal seed coat with a tracheoidal exotegmen. The tribe Dicoelieae is also distinct from the other Acalyphoideae in having an exotegmic seed that is composed of ribbon-like cells of exotegmen (i.e., cells both longitudinally and radially elongated, sclerotic and pitted). The tribe Galearieae, which should be treated as a distinct family Pandaceae, is also distinct from the other Acalyphoideae in having an exotegmic seed with a tracheoidal exotegmen (i.e., cells longitudinally elongated, sclerotic and pitted). The remaining genera of Acalyphoideae always have an exotegmic seed with a palisadal exotegmen (i.e., cells radially elongated, sclerotic and pitted). The shared palisadal exotegmen supports the close affinity of Acalyphoideae (excluding five tribes) with Crotonoideae and Euphorbioideae. Within the remaining genera of Acalyphoideae, a significant diversity is found in ovule and seed morphology with respect to the thickness of the inner and outer integument, the size of chalaza, vascularization of an outer integument and an aril.     

Seed coat morphology and its systematic significance in Juncus L. (Juncaceae) in Egypt

The seed morphology of nine taxa of Juncus from Egypt has been investigated using light and scanning electron microscopy, to determine the importance of seed coat features as taxonomic characters. Macro- and micromorphological characters, including seed shape, color, size, seed appendages, epidermal cell shape, anticlinal boundaries, and outer periclinal cell wall and secondary cell wall sculpture are presented. Four types of seed appendages are recognized: (i) seeds with two appendages; (ii) seeds without appendages; (iii) seeds with minutely a piculate at one end; and (iv) seeds with minutely a piculate at both ends. Two types of anticlinal cell wall boundaries,(i) raised-channeled, straight and (ii) raised, straight or sinuous, and three different shapes of outer periclinal cell wall are described: (i) flat; (ii) concave; and (iii) flat to slightly concave. The secondary sculpture of the cell wall varies from striate to microreticulate or reticulate, and smooth to finely folded. Seed characters provide useful data for formulating the taxonomy of Juncus both on the subgeneric and sectional level. A key for the identification of the investigated taxa based on seed characters is provided.     

Seed coat morphology and its systematic significance in Juncus L. (Juncaceae)in Egypt

The seed morphology of nine taxa of Juncus from Egypt has been investigated using light and scanning electron microscopy,to determine the importance of seed coat features as taxonomic characters.Macro-and micromorphological characters,including seed shape,color,size,seed appendages,epidermal cell shape,anticlinal boundaries,and outer periclinal cell wall and secondary cell wall sculpture are presented.Four types of seed appendages are recognized:(i)seeds with two appendages;(ii)seeds without appendages;(iii)seeds with minutely a piculate at one end;and(iv)seeds with minutely a piculate at both ends.Two types of anticlinal cell wall boundaries,(i)raised-channeled,straight and(ii)raised,straight or sinuous,and three different shapes of outer periclinal cell wall are described:(i)flat;(ii)concave;and(iii)flat to slightly concave.The secondary sculpture of the cell wall varies from striate to microreticulate or reticulate,and smooth to finely folded.Seed characters provide useful data for formulating the taxonomy of Juncus both on the subgeneric and sectional level.A key for the identification of the investigated taxa based on seed characters is provided.     

Taxonomy of the Genus Oreocharis (Gesneriaceae)

The genus Oreocharis as circumscribed here consists of 27 species including 5 varieties, of which 5 species and 4 varieties are described as new in the present paper. In the work analysed were the external morphology and geographic distribution and examined under SEM were pollen exine of 22 species and seed coat of 16 species. As a result, three types of the corolla, two types of the anther, three types of the pollen exine and three types of the seed coat are distinguished here in the paper. It is discovered that the corolla in the genus is relatively stable, though diverse, and highly correlated with the characters of pollen grains and seeds. The corolla clearly bilabiate but constricted at the throat, occurring in O. auricula, O. cordatula, O. aurantiaca, etc., for an example, is correlated with smooth, reticulate pollen exine and partial tectum and the reticulate and smooth seed coat. For this reason the subdivision of the genus in the paper is mainly based on the characters of the corolla, but combined with those of the anther, pollen and seed coat. The genus is divided into four sections in the present classification. Dasydesmus Craib, based on a single species. O. bodinieri, is reduced here, and the reasons are given. The genus is distributed mainly in the subtropics, and less frequently in the tropics, of China south of 32.5°N and east of 98.5°E, with only two species beyond the border, O. hirsuta in Thailand (only a single locality in Chiengmai) and O. aurea also found in north Vietnam (see Fig. 1, Table 3). Sect. 1. Stomactin (Clarke) Fritsch. Corolla urceolate-tubular, constricted at the throat, with limb distinctly bilabiate; anthers broad-oblong; seed coat reticulate, smooth, rarely minutely tuberculate; pollen exine fine-reticulate, tectum partial and smooth, luminae slightly unequal in size. Sect. 2. Orthanthera K. Y. Pan Corolla campanulate or campanulate-tubular; anthers broad-oblong; seed coat reticulate, muri smooth, rarely spiny-processed; pollen exine fine-reticulate, with partial and smooth tectum and luminae slightly unequal in size, rarely exine insular and fine-tuberculate, tectum perforate. Setc. 3. Oreocharis Corolla thin-tubular; anthers broad-oblong; seed coat densely spinyprocessed, rarely fine-tuberculate; pollen exine insular, densely spiny-processed, rarely finereticulate and smooth, luminae unequal in size. Sect. 4. Platyanthera K. Y. Pan Corolla campanulate; anthers hippocrepiform; seed coat densely spiny-processed; pollen exine fine-reticulate, tectum perforate, luminae small, nearly equal in size. In the section Stomactin, although the constriction of corolla at its throat is a specialized character, the characters of seed coat, pollen grains and anthers are apparently primitive. Therefore it may be said at least that more primitive characters are preserved in the section. In the section Oreocharis, on the contrary, the characters of corolla, seed coat and pollen exine are all advanced. And in the section Platyanthera, the seed coat, pollen (with perforate tectum) and anthers have developed rather specialized characters.