Pollen morphology of the tribe Cynoglosseae of Boraginoideae (Boraginaceae) in China

Pollen morphology of 16 Chinese species representing 7 genera in the tribe Cynoglosseae of Boraginoideae(Boraginaceae) was examined under LM and SEM, and 5 species under TEM. Pollen grains are cocoon-shaped, rarely subprolate, prolate or ovoid, very small, 7～ 15.7μm× 3.5 ~ 13.9 μm in size, P/E = 1.6～2.02; 3-colporate apertures alternate with 3-pseudocolpi, with equatorial endocingulus except those in Bothriospermum; exine surface is usually smooth, with or without perforations in two poles, rarely with tuberculate ornamentation; exine is rather thin and includes ectexine and endexine, while ectexine consists of imperforate tectum, columellae and foot-layer. The tribe Cynoglosseae has many common characters of pollen grains, but there are some differences among genera. A key to the genera is given based on pollen morphology. Bothriospermum, without endocingulus, may be a primitive genus in this tribe, and its pollen morphology is more similar to that of Eritrichieae than to Cynoglosseae, thus it seems more reasonable to put it into Eritrichieae. Solenanthus may be the most advanced genus in the tribe Cynoglosseae for its pollen grains of sub-isopolar and ovoid shape.     

A phylogenetic analysis of morphological and molecular characters of Boraginaceae: evolutionary relationships,taxonomy, and patterns of character evolution

The angiosperm family Boraginaceae includes ca. 1600 species distributed among ca. 110 genera. Some floral features are constant within the family, but many vegetative, floral, pollen, and nutlet traits vary. Utilizing 224 species of Boraginaceae and related taxa, five matrices were constructed with various combinations of morphological characters, three chloroplast DNA regions, and one nuclear ribosomal DNA region. Phylogenetic analyses were conducted for these matrices, and patterns of character evolution were examined. Boraginaceae is resolved as monophyletic, with Wellstedia as its sister. Codon is sister to Boraginaceae + Wellstedia. Although most of the investigated morphological characters have a low consistency index, particular character states are synapomorphies for large clades in each of the tribes of the family. In Boraginaceae, the breeding system heterostyly evolved at least 12 times, which is the largest number of origins resolved in any family; therefore Boraginaceae can serve as a model for the evolution and development of heterostyly. Nutlet ornamentation is most diverse in Cynoglosseae and Trichodesmeae, while pollen and floral features are most variable in Boragineae and Lithospermeae. Phylogenetic relationships and patterns of character evolution identified in the present study set the stage for future work creating an updated taxonomic system of Boraginaceae.     

Karyological studies on Spanish Boraginaceae. VI. Contribution to the tribe Eritrichieae

LUQUE, T., 1992. Karyological studies on Spanish Boraginaceae. VI. Contribution to the tribe Eritrichieae. Karyological information is presented for c. 15 taxa of the genera Asperugo L., Rochelia Reichenb., Myosotis L. and Lappula Gilib. This paper is the first in this series of karyological studies of Spanish Boraginaceae to deal with Eritrichieae.     

Tribe Eritrichieae (Boraginaceae s.str.) in West Asia: a molecular phylogenetic perspective

A selection of Boraginaceae genera was used to obtain a framework for the phylogenetic position of some tribes belong to subfamily Boraginoideae and genera within tribe Eritrichieae (Heterocaryum, Rochelia, Eritrichium, Lappula, Lepechiniella, and Asperugo) and related species. Our results were produced on the basis of nrDNA ITS and cpDNAtrnL-F sequences. The combined nrDNA ITS trnL-F data confirm four main clades of Boraginoideae comprising Echiochileae, Boragineae, Lithospermeae, and Cynoglosseae s. l. (including Eritrichieae, Cynoglosseae s. str., and Myosotideae). The tribe Eritrichieae itself at the current status is paraphyletic; some members, for example Asperugo procumbens, Lepechiniella inconspicua, Myosotidium hortensia, and Cryptantha flavoculata are placed out of the core tribe Eritrichieae. The genus Heterocaryum is monophyletic and allied with a subclade of genera Lappula, Lepechiniella, Eritrichium, and Rochelia. Rochelia is monophyletic, but Eritrichium and Lappula are non-monophyletic. Lepechiniella is nested among a group of Lappula species.     

Tribes of Boraginoideae (Boraginaceae) and placement of Antiphytum, Echiochilon, Ogastemma and Sericostoma: A phylogenetic analysis based on atpB plastid DNA sequence data

 The genera Antiphytum, Echiochilon, Ogastemma and Sericostoma, which have been difficult to place within the tribes of the subfamily Boraginoideae (Boraginaceae), are analysed using plastid atpB sequence data. A selection of Boraginaceae genera was used to obtain a framework for the phylogenetic position of Antiphytum, Echiochilon, Ogastemma and Sericostoma. Sericostoma is found to belong within Echiochilon. The new tribe Echiochileae, Boragineae and Lithospermeae are monophyletic but the tribes Eritrichieae and Cynoglosseae are paraphyletic. The biogeography of Echiochileae (Echiochilon and Ogastemma from Africa and western Asia, and Antiphytum from America) is discussed. Received November 27, 2001; accepted March 20, 2002 Published online: November 14, 2002 ? Springer-Verlag 2002 Addresses of the authors: Elisabeth L?ngstr?m (e-mail: elisabeth.langstrom@ebc.uu.se), Department of Systematic Botany, Evolutionary Biology Centre, Uppsala University, Norbyv?gen 18D, SE-752 36 Uppsala, Sweden. Mark W. Chase, Jodrell Laboratory, Royal Botanic Gardens, Kew, TW9 3DS, UK.     

Pollen morphology of the tribe Lithospermeae of Boraginoideae in China and its taxonomic significance

The pollen morphology in 15 species representing five genera in the tribe Lithospermeae of Boraginoideae (Boraginaceae) has been investigated and illustrated using light microscopy and scanning electron microscopy. The tribe Lithospermeae is very diverse in pollen morphology. The pollen grains are 10.4–41.8 × 7–33.1 μm in size with subspheroidal, prolate, cocoon, dumbbell, and ovoid shapes. The pollen apertures are of five types: 3-colporate, 3-syncolporate, 4-8-colporate, 4-6-syncolpate, and 6-7-colpate types. The exine ornamentations are generally smooth or rugulose, sparsely echinulate, and rarely rugulate. Based on pollen morphology, we developed a key to identify the genera of the tribe Lithospermeae, compared the pollen apertures among tribes of Boraginoideae, explained the evolutionary trends of the pollen grains, and discussed the taxonomic position of the tribe Lithospermeae. The palynological data suggest pollen of the eurypalynous type and support the proposal that the tribe Lithospermeae is in the primitive position of the subfamily Boraginoideae, and Echium Linn. is in the tribe Lithospermeae rather than in a novel tribe. Our observations have application potential for identification of pollen fossils of the tribe Lithospermeae.     

Taxonomy and distribution of the genus Rubus (Rosaceae) series Radula on the Iberian Peninsula

Monasterio-Huelin. E. 1995. Taxonomv and distribution of the genus Rubus (Rosaceae) series Radulu on the Iberian Peninsula. — Nord. J. Bot. 15: 365–373. Copenhagen. ISSN 0107–055X.
The majority of european batologists currently interpret the ser. Radula in a sense different from that given by its original author. According to the modem concept, this series is separated into two, distinguished by the leaflets felted beneath (ser. Radula ) versus not felted (ser. Pallidi ). Following this criterion, we include 7 species in ser. Radula , one of which ( R. pauanus ) is described as new, and provide a photograph of the holotype, an analysis of pollen, and a distribution map. Also provided are keys to the series of the subgenus Rubus and to the species of the ser. Radula , and a comparative study from a morphological and biogeographical perspective of all species which comprise this series on the Iberian Peninsula.     

Stigma characteristics and angiosperm taxonomy

When a classification based upon certain morphological and physiological characters of the stigma was applied to ca. 1,000 species of some 900 genera of angiosperms, several taxonomic regularities emerged. Most families proved to be relatively homogeneous in stigma type, whilst others were strikingly diverse. Among the latter were certain supposedly primitive families such as the Liliaceae, a fact that may have some phylogenetical significance. Sometimes where a family proved to be homogeneous in respect to the more general characters of the stigma, fuller investigation revealed great variation in detail. An example is given by the Boraginaceae, a family in which stigma characters are of potential taxonomic importance. The stigma papillae in this family are often capitate, with heavily cutinised heads, and pollen has to be inserted forcibly between the papillae. There is a strong correlation between pollen size and stigma–papilla size, so that certain types of cross pollination are impossible. These specialisations can thus play a part as isolating mechanisms. The form of the stigma papillae is probably also important in genera of the Boraginaceae adapted to arid conditions, since the contiguous cutinised caps overarch the actual receptive surfaces and so. presumably, offer protection from desiccation.
In both monocotyledons and dicotyledons, clear relationships exist between stigma type and physiology, and notably with the type of self–incompatibility system. It is already apparent that stigma characters can have predictive value in the investigation of breeding systems.
The paper includes a comprehensive literature list, covering most ot the principal sources of structural, taxonomic and other data bearing upon angiosperm stigmas.     

New light into the hormogastrid riddle: morphological and molecular description of Hormogaster joseantonioi sp. n. (Annelida,Clitellata, Hormogastridae)

The earthworm family Hormogastridae shows a remarkable disjunction in its distribution in the Iberian Peninsula, with the Hormogaster elisae species complex isolated from the rest of the species. Hormogaster joseantonioi sp. n., a new species found in the intermediate area between the main ranges (in Teruel, Aragón), was described following the integrative approach, as it is suitable for earthworms due to their highly homoplasic morphology. The phylogenetic analysis of the molecular markers placed the new species as a sister taxon to H. elisae, thus showing the colonizing lineage of Central Iberian Peninsula could have originated near the H. joseantonioi sp. n. current range. External morphological characters revealed some degree of overlap with previously described species, but internal characters presented configurations/states unknown from other members of the family. These traits make the new species a key piece to understand the evolution of Hormogastridae.     

Predictive modelling of tree species distributions on the Iberian Peninsula during the Last Glacial Maximum and Mid-Holocene

This paper reports a bioclimatic envelope model study of the potential distribution of 19 tree species in the Iberian Peninsula during the Last Glacial Maximum (LGM; 21 000 yr BP) and the Mid-Holocene (6000 yr BP). Current patterns of tree species richness and distributions are believed to have been strongly influenced by the climate during these periods. The modelling employed novel machine learning techniques, and its accuracy was evaluated using a threshold-independent method. Two atmospheric general circulation models, UGAMP and ECHAM3 (generated by the Palaeoclimate Modelling Intercomparison Project, PMIP), were used to provide climate scenarios under which the distributions of the 19 tree species were modelled. The results obtained for these scenarios were assessed by agreement measure analysis; they differed significantly for the LGM, but were more similar for the Mid-Holocene.
The results for the LGM support the inferred importance of pines in the Iberian Peninsula at this time, and the presence of evergreen Quercus in the south. Important differences in the altitude at which the modelled species grew were also predicted. During the LGM, some normally higher mountain species potentially became re-established in the foothills of the Pyrenees. The warm Mid-Holocene climate is clearly reflected in the predicted expansion of broad-leaved forests during this period, including the colonization of the northern part of the Iberian Peninsula by evergreen Quercus species.     

A new species of Hackelia (Boraginaceae) from Utah

Hackelia ibapensis, a new species of Boraginaceae (Subfamily Boraginoideae, Tribe Eritrichieae) from the Deep Creek Range of Utah is described and illustrated.     

Pollen of African Spermacoce species (Rubiaceae) Morphology and evolutionary aspects

Pollen morphology of 43 African species of the genus Spermacoce has been investigated by scanning electron and light microscopy. The genus is eurypalynous, which is reflected in the remarkable variation of almost all pollen characters. The average equatorial diameter (E) ranges from 15.8 w m to 115.5 w m. Grains are colporate or pororate. The number of apertures varies from 3 up to more than 25. The majority of species has apertures situated only at the equator (being zonoaperturate), but a few species have pantoaperturate grains. The endoaperture is generally an endocingulum, often with a secondary lolongate or lalongate thinning at the ectocolpus; endocolpi and endopores are also observed. The sexine is usually perforate, but eutectate, foveolate, and (micro)reticulate tecta were also found. Supratectal elements are present as granules, microspines or spines. The inner nexine surface is granular, often with irregular grooves (endocracks). Among native African species, nine pollen types are recognized mainly on the basis of pollen size, aperture morphology and tectum peculiarities. In two of the pantoaperturate types, apertures are in a configuration not yet recorded for the angiosperms in general. Some evolutionary trends are proposed that await verification by further systematic study. Pollen morphological characters have a high taxonomic value in the genus Spermacoce . They provide almost unique identification marks for the species, which enables sharpening of species boundaries. Small groups of related species often share the same pollen type. The genus Borreria , previously separated from Spermacoce on the basis of its fruit morphology only, is not supported by pollen data.     

Pollen Morphology and Epidermal Characters of Leaves in Convallarieae (s. l.)

The tribe Convallarieae (sensu Krause 1930) consists of 7 genera, i.e. Convallaria, Speirantha, Reineckia, Theropogon, Tupistra, Rohdea and Aspidistra, but now generally recognized as two tribes, Convallarieae (the former 4 genera) and Aspidistreae (the rest). Observed in this work were pollen morphology of 17 species and epidermal characters of leaves of 12 species. All the 7 genera are covered in observations. Pollen grains in Convallarieae (s. str.) are all monosulcate and boat-shaped (Plate 1: A-F). The exine is rather uniformly microperforate (Plate 1: A-F); only Theropogon is exceptional in this respect: it has rugulate exine (Plate 1: O, P). Tang and Zhang (1985) have pointed out the heterogeneity of Theropogon in this tribe. Pollen morphology in the tribe Aspidistreae is widely variable. The genera Tupistra and Rohdea were shown to have monosulcate and boat-shaped pollen grains. Their exine is perforate or reticulate (Plate 1: G-N). Pollen grains in the genus Aspidistra, however, are nonaperturate and spheroidal. The exine in the genus varies from crass-rugulate, variously gemmate to tuberculate-baculate (Plate 2; A-H). The pollen morphology of Aspidistra is therefore distinctly different from that of Tupistra and Rohdea, which supports the Nakai's (1936) establishment of the tribe Rohdeae for Tupistra and Rohdea. Therefore, Krause's Convallarieae is reasonably divided into at least three tribes, Convallarieae (Speirantha, Convallaria, Reineckia and Theropogon), Aspidistreae (Aspidistra) and Rohdeae (Rohdea and Tupistra). The pollen characters of all the 7 genera are shown in Table 1. The evolutionary trends of pollen morphology (aperture and exine) in the three tribes are discussed and our major view-points are shown in Fig. 1. Observations on epidermal characters of leaves show that in the Convallarieae (s. 1.) stomatal apparatuses are all anomocytic; cuticular layer on the upper epidermis is mainly striatethickened or rather uniformly thickened (Plate 2: J--P; Plate 3: A-C, F-N), whereas in the genus Convallaria the cuticular layer is squamosely thickened (Plate 2: I; Plate 3: D, E).The epidermal characters of leaves in the 7 genera are summarized in Table 2.     

中国紫草科厚壳树亚科的花粉形态及其系统学意义

为探讨厚壳树亚科在紫草科中的系统位置,用扫描电子显微镜研究了国产厚壳树亚科Ehretioideae的厚壳树属Ehretia、轮冠木属Rotula和基及树属 Carmona 3属12种植物的花粉形态,其扫描电镜资料属首次报道。从本文并结合文献报道的双柱紫草属 Coldenia的花粉资料,可得知该亚科花粉为长球形 、近球形,稀球形,赤道部位具有不同程度的膨大;萌发孔类型为三孔沟和三拟假沟二者交替排列,稀具三孔;外壁表面具微弱的小穿孔,稀光滑或皱波状纹饰。厚壳树亚科Ehretioideae与破布木亚科     

Pollen diameter and fertility in nine species ofPuccinellia (Poaceae)

Diameter and fertility of pollen grains were surveyed in nine species ofPuccinellia Parl. (Poaceae), mainly from the Iberian Peninsula, allowing information about reproductive behaviour and biological characteristics of pollen of these taxa. Maximum diameter was measured with an image processor and fertility was evaluated by staining the grains with Müntzing's mixture. Significant differences in both characters were found between the taxa. In consequence, maximum diameter and fertility proved to have some taxonomic relevance inPuccinellia.     

Sexual reproduction in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The reproductive biology of someScrophularia species in the Iberian Peninsula and Balearic Islands has been studied. Levels of fruit-set in bagged and unbagged inflorescences, the production of seeds/capsule, the effectiveness of seed-set following cross-pollination in the genus and the pollen/ovule ratios have been analyzed.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and Balearic Islands. II. For first part seeOrtega Olivencia & Devesa 1993.     

中国紫草科破布木属花粉形态和外壁超微结构

为了深入探讨紫草科（Boraginaceae）的分类问题,用光学显微镜和扫描电子显微镜观察了该科破布木属（CordiaL.) 10种植物的花粉形态和外壁超微结构。发现该属花粉具三孔、三孔沟、三拟孔沟和三合沟4种萌发孔类型。外壁表面具微刺状纹饰、刺状纹饰、网状纹饰和不规则的条纹网状纹状。破布木属的花粉特征表明,该属花粉在紫草科中既是独特的分类群,又是比较原始的属种。     

Some considerations about the postglacial history and recent distribution ofFagus sylvatica in the NW Iberian Peninsula

This paper presents a re-assessment of the evidence of beech (Fagus sylvatica) presence on the NW Iberian Peninsula in the different stages of the Late Quaternary. For this purpose, numerous pollen diagrams were re-examined and the present-day distribution of the species was documented. Beech pollen has been recorded in different periods, before the Würm Pleni-Glacial, during the Late Glacial and the Holocene, leading to the hypothesis that there existed local refugia forFagus sylvatica in the NW Iberian Peninsula, comparable to those in more eastern areas of southern Europe, the existence of which has been recognized.     

Pollen Morphology and Wall Stratification in Rubus L. (Rosaceae) in the Iberian Peninsula

A palynological study using LM and SEM was made of the morphology, ornamentation, apertural system and structure of the pollen wall, of the 18 species of the genus Rubus which are present in the Iberian Peninsula. We demonstrate: a) the stenopalynous character of the genus (isopolar, subprolate, 3-lobate, 3-zonocolporate and striate-perforate, except for R. genevierii whose ornamentation is striate-gemmate-perforate and with a striated margin); b) the lalongate shape of the endoaperture, due to the disappearance or thinning of the nexine; and c) the existence of two structural types, Type I granular and Type II columellar with clearly-defined columellar spaces or compact without intercolumellar spaces.     

Floral rewards in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The genusScrophularia in the Iberian Peninsula and Balearic Islands comprises two sections,Scrophularia andCanina G. Don. Analyses were carried out on flower production, flower duration and their sexual phases, pollen and nectar production together with observations on their pollinators. Nectar production is correlated with corolla size and pollen production with anther size. The taxa of sect.Scrophularia show greater nectar and pollen production than those of sect.Canina. Also, those of the first section produce more ovules per ovary than those of the second group, production being correlated with the ovary size.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and the Balearic Islands. I.