From acaveate to caveate: evolution of pollen types in the Rhaponticum group (Asteraceae,Centaureinae) related to extreme conditions

A survey of pollen morphology and ultrastructure within the Rhaponticum group, using both scanning and transmission electron microscopy, revealed two pollen types. Most representatives of the Rhaponticum group exhibited Serratula‐type pollen. The exceptions were the genus Oligochaeta, with Centaurea jacea‐type pollen, and the genus Myopordon, with both types of pollen. Phylogenetic inference within the group indicated that Jacea‐type pollen is derived from Serratula‐type pollen. The fact that Myopordon exhibited a series of pollen types, from Serratula type to Jacea type, suggests that a transition from one form to another has occurred in the genus. Therefore, the factor which has produced this evolutionary tendency should be sought within Myopordon. Jacea‐type pollen is characterized by a decrease in size, ornamentation, the absence of a columellar infratectum and the presence of a cavea. A comparative survey of similar tendencies in other groups, and their correlation with potential functional implications, prompted the hypothesis that the critical factor involved in such evolution of pollen could be habitats with extreme conditions. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 499–510.     

SPORE MORPHOLOGY IN THE GENUS BRUCHIA SCHWAEGR. (MUSCI)

Scanning electron microscopy observations of the spores of Bruchia have resulted in the recognition of four spore types based on the ornamentation of the distal spore surface: warty or verrucate, pitted, reticulate, and spinose. The proximal surface of the spores of all species, except B. brevipes, is characterized by a central aperture region surrounded by a triangular murus or rows of spinae forming a triangle and 1 or 2 smooth or verrucate collars. The ornamentation patterns observed are considered to be characteristic for the genus. Spore morphology alone can rarely be used to distinguish species but in conjunction with certain other characteristics, it is an important taxonomic feature. Spore morphology is a major characteristic used to define the limits of the highly variable species B. flexuosa (spinose spores) and to distinguish it from the closely related species B. texana (reticulate spores). Variations in ornamentation patterns within the genus support the recognition of two subgenera (subgenus Bruchia and subgenus Sporledera). Spore morphology also supports the close relationship between Bruchia and Trematodon and is sufficient to eliminate several questionable taxa from the genus.     

Fruit and seed morphology of the tribe Orobancheae (Orobanchaceae) genera in Turkey and its taxonomic significance

Seed and fruit morphology of four genera of Orobanchaceae: Cistanche, Diphelypaea, Orobanche and Phelipanche, native to Turkey, were studied using light and scanning electron microscopy. In this study, 100 accessions belonging to 39 species were investigated. The examined fruits had loculicidal capsules, generally dehiscing with 2 slits, and the seeds were small and with reticulate ornamentation. The basic shape of the seeds in most of the studied taxa was oblong, ellipsoid to ovoid, globose or subglobose. The results indicate that characters such as seed size, shape and color are useless for identification of the taxa at the species level. However, the anticlinal wall pattern and periclinal wall ornamentation proved useful for identifying the different genera. Four morphological types could be distinguished: smooth, perforate, microfibrillar and granulate. Type I corresponds to the genus Cistanche and Orobanche anatolica, whereas types II, III and IV were found in Diphelypaea, Orobanche and Phelipanche, respectively. The utility of seed characteristics is discussed in terms of inter‐ and intra‐generic taxonomy.     

Pollen diversity in the genus Ptilostemon (Asteraceae,Cardueae) from Italy and its taxonomic and palynoecological implications

The results of a study on pollen morphology of the species of the genus Ptilostemon growing in Italy: Ptilostemon niveus, P. greuteri, P. gnaphaloides, P. casabonae, P. strictus, P. stellatus by light and scanning electron microscopy were presented and discussed including in relation to their taxonomic position. The exine shows two different ornamentation patterns: echinae and scabrae. This last ornamentation pattern together with other features was previously described in the genus Ptilostemon. Of 6 species analysed, P. stellatus is the only one that shows a scabrate ornamentation of exine. Two pollen types were recognized through the exine ornamentation, length of ectocolpus and polar outline: P. stellatus type and P. niveus type. A multivariate analysis (principal component analysis and linear discriminant analysis) was carried out with the aim of examining potential morphological characters which could be used to identify taxa. The data suggest that several characteristics can be used to delimit the species in particular the exine ornamentation. The results of our studies support the actually subgeneric classification, and several features of the pollen grains analysed seem to have a palynoecological role and an important taxonomic significance. A dichotomous key based on palynological data is also given.     

Pollen morphology of Alcantarea giant bromeliads (Bromeliaceae,Tillandsioideae)

Here we describe the pollen morphology of 19 species of Alcantarea, none of them fully described before, to contribute to the infra-generic delimitation of the genus. Pollen grains are heteropolar, amb ellipsoidal (although spheroidal pollen occurs in Alcantarea aurantiaca, A. burle-marxii, A. glaziouana, A. nahoumii, A. roberto-kauskyi and A. trepida), monosulcate, sulcus with margin, reticulate ornamentation of the exine on the central area of pollen grain (except in A. imperialis, which display foveolate ornamentation) and apex of equatorial axe with different ornamentation from the central area of pollen grain (microreticulate calota or psilate-perforate calota). The genus is considered stenopalynous due to the similarity in the pollen morphology. The new pollen characters obtained here are useful in understanding the infra-generic palynotaxonomy of the genus confirming the relationship between species that have been grouped together based on vegetative evidence and/or relationships suggested by analyses based on DNA sequences or microsatellite evidence, e.g. between A. aurantiaca, A. burle-marxii and A. nahoumii; between A. burle-marxii, A. glaziouana and A. trepida; between A. hatschbachii and A. duarteana, between A. roberto-kautsky and A. trepida. Our results improve the morphological characterisation of the genus and may contribute to phylogenetic analyses as well as the biosystematics approach to solve species complexes.     

Systematic implications of pollen morphology in Elsholtzia (Elsholtzieae–Lamiaceae)

Pollen morphology of 18 species (32 specimens) representing all three currently recognized sections of the genus Elsholtzia (Elsholtzieae–Lamiaceae) was investigated in detail using light, scanning electron and transmission electron microscopy. Elsholtzia pollen grains are small to medium in size (P=20–50 μm, E=16–45 μm), mostly prolate‐spheroidal to prolate, and rarely oblate‐spheroidal to subprolate in shape (P/E=0.95–1.46), hexacolpate (the amb more or less circular or ellipsoid) with granular aperture membranes. Three distinct types of sexine ornamentation are observed in Elsholtzia: perforate, rugulose‐bireticulate, and bireticulate. The bireticulate pattern is the most common, showing a wide range of morphological variation among species. The exine thickness varies from 1.2 to 2.5 μm. Among the present palynological data, the variability observed in sexine ornamentation of Elsholtzia could be of systematic significance. Current infrageneric classification of the genus Elsholtzia is partially congruent with pollen morphological data.     

NOTES ON SOME GRASSES. XVI. EMBRYO STRUCTURE OF THE GENUS ORYZA IN RELATION TO THE SYSTEMATICS

Three distinct types of embryo have been found in 18 species of the genus Oryza. Type 1 is characterized by lateral fusion of the epiblast and the scutellum and also by the development of auricles. Type 2 has no auricle, and the epiblast is not laterally fused with the scutellum. Type 3 has a cleft between the lower part of the scutellum and the coleorhiza, a long epiblast which is free from the scutellum, and lacks auricles. Type 1 has been found in 13 species of section Sativae, Type 2 in 4 species of sections Granulatae and Coarctatae, and Type 3 in O. coarctata only. The results indicate that species of section Sativae are more or less closely related, while section Coarctatae is a heterogeneous group. A new section, Ridleyianae, has been described for Oryza ridleyi and some other species. Oryza coarctata has unique features in embryo structure, morphology and leaf anatomy, and this species has been moved from the genus Oryza to a separate genus, Sclerophyllum, which is here revived.     

An electron microscopical study of the genus Tortula Hedw., with special reference to exine ornamentation

Abstract

Spores of ten North European species of the genus Tortula Hedw. were studied in the transmission electron microscope. In all the sporoderm was more or less covered with finely papillate processes of several different types which divide the genus into a number of groups different from the previously suggested sections. The ornamentation of leaves, peristomes, and spores of T. norvegica (Web. f.) Wahlenb. has been studied in the scanning electron microscope. The leaf papillae are not c-shaped as suggested in the literature, but branched in a rather complex way. The borders of the basal membrane cells of the peristome appeared to be a continuation of the filamentous portion of the teeth. A comparison with T. ruralis (Hedw.) Gaertn., Meyer & Scherb. revealed the ornamentation to be of the same basic type in the two species.     

A new Patagonian species of Cricosaurus (Crocodyliformes,Thalattosuchia): first evidence of Cricosaurus in Middle–Upper Tithonian lithographic limestone from Gondwana

Abstract:  Upper Jurassic (Tithonian) deposits in north‐western Patagonia, Argentina, have yielded rich and taxonomically diverse assemblages of marine reptiles. These assemblages are also remarkable by their quality of preservation and are represented by ichthyosaurs, plesiosaurs, turtles and crocodyliforms. Despite the abundant crocodyliform record, only two metriorhynchid taxa have been identified: Cricosaurus araucanensis and Dakosaurus andiniensis. Here we described a new species of Cricosaurus, which represents the second species of Cricosaurus in the Tithonian of the Neuquén Basin, and the first metriorhynchid found in lithographic limestone from Gondwana. Furthermore, this specimen has the most complete postcranial remains of any metriorhynchid from South America. The new species is characterized by a short distance between the premaxilla and the nasal, a relatively narrow interorbital width, 23–25 mandibular teeth, bicarinated teeth with fine apicobasally aligned ridges, interalveolar spaces between the first seven teeth approximately 1.5 times longer than the anteroposterior diameter of the respective alveoli. To test the assignment of the new species to Cricosaurus, we carried out two phylogenetic analyses. In both analyses, Cricosaurus lithographicus sp. nov. is nested with other species referred to this genus. This new species has peculiar enamel ornamentation, characterized by numerous, fine apicobasally aligned ridges, when compared to other species of the genus.     

Mychophilus palmatus, a new species (Copepoda: Cyclopoida: Ascidicolidae) associated with the compound ascidian Botryllus leachi Savigny, with remarks on congeneric species and related genera

Presently two species of the genus Mychophilus Hesse, 1865 are known: Mychophilus roseus Hesse, 1865 and Mychophilus fallax Stock, 1967. A hitherto unknown species is described herein as Microphilus palmatus n. sp. The new species is characterized by its urosome much exceeding the cephalosome-metasome in length, the ornamentation of the labrum, the shape of the palp of the first maxilla, the ornamentation of the legs 1–4 and shape of the caudal rami. These and other features of the genus Mychophilus are compared with those from other related genera of the family Ascidicolidae Thorell, 1859.     

Pollen morphology and infra-generic evolutionary relationships in some Chinese species of Pedicularis (Scrophulariaceae)

 The pollen grains of 32 Chinese species of Pedicularis representing 13 groups within the genus were investigated using light microscopy and scanning electron microscopy. The genus is eurypalynous, the apertures may be tricolpate, trisyncolpate, or bisyncolpate and five types of surface ornamentation (microscabrate, microrugulate, retipilate, microreticulate and microfoveolate) were observed. Comparison of the different aperture types using light microscopy allows three pollen types to be distinguished. Examination of exine ornamentation with scanning electron microscopy enabled each pollen type to be divided into two or three sub-types (giving a total of seven sub-types). Pollen morphology within the genus is discussed in relation to infrageneric relationships, evolutionary and pollination biology. There is little correlation with the existing infrageneric taxonomy (some taxonomic groups have more than one pollen type, while the same pollen type may be found in several different species-groups), but greater correlation with the corolla morphology. Received January 10, 2002; accepted March 5, 2002 Published online: March 10, 2003     

Studies in Heterobasidiomycetes,Part 38*)Sphacelotheca polygoni-persicariae G. Demi & Oberw. spec. nov.

A new species, Sphacelotheca polygoni-persicariae, parasitizing the ovaries of Polygonum persicaria L. is described. This smut, collected several times in Madeira, Portugal, differs from other species of that genus, by its host plant and the reticulated teliospore ornamentation. On the basis of the morphological and ultrastructural characters of S. polygoni-persicariae, in connection with some recently published data on siderophore formation and 5S ribosomal RNA sequences it can be assumed that 1) members of the genus Sphacelotheca are separated from typical Ustilago species of Poaceae, 2) Sphacelotheca is restricted to species parasitizing Polygonaceae, and 3) species of Sphacelotheca and Microhotryum as well as those Ustilago species which parasitize in the flowers of Polygonaceae are closely related.     

Osservazioni preliminari al microscopio elettronico a scansione sulle ornamentazioni delle spore di Russula

Abstract

Preliminary observations on spore ornementation of Russula, as seen in the scanning electron microscope. — The spores of 16 species of Russula have been examined in the scanning electron microscope, as a preliminary attempt to see if an accurate examination of the spore surface at the ultrastructural level could reveal details of ornamentation which might be useful for the classification of the many species of this genus. The examination, carried out both on fixed and unfixed specimens, has demonstrated that the spore shape is always round or slightly elliptic, and that the obnormal forms as previously described are probably artifacts. Five main types of ornamentation have been described: single wart-like and single finger-like Protrusion, wart- or finger-like protrusions interconnected by thin ridges, and thick, short « papillae » together with large ridges that run along large tracts of the spore circumference. The type of ornamentation was a constant character in each species.     

New Striate Pollen from the Pennsylvanian of Kansas

A new genus of striate saccate pollen, Caelopollis is described from the Virgilian (Pennsylvanian) of Kansas. A striate distal leptoma serves to distinguish the genus and four new species of striate pollen. One new species of Vittatina, V. moorei, is described that also possesses an unusual type of distal ornamentation.     

Testing the success of palaeontological methods in the delimitation of clam shrimp (Crustacea,Branchiopoda) on extant species

Fossil spinicaudatan taxonomy heavily relies on carapace features (size, shape, ornamentation) and palaeontologists have greatly refined methods to study and describe carapace variability. Whether carapace features alone are sufficient for distinguishing between species of a single genus has remained untested. In our study, we tested common palaeontological methods on 481 individuals of the extant Australian genus Ozestheria that have been previously assigned to ten species based on genetic analysis. All species are morphologically distinct based on geometric morphometrics (p ≤ 0.001), but they occupy overlapping regions in Ozestheria morphospace. Linear discriminant analysis of Fourier shape coefficients reaches a mean model performance of 93.8% correctly classified individuals over all possible 45 pairwise species comparisons. This can be further increased by combining the size and shape datasets. Nine of the ten examined species are clearly sexually dimorphic but male and female morphologies strongly overlap within species with little influence on model performance. Ornamentation is commonly species-diagnostic; seven ornamentation types are distinguished of which six are species-specific while one is shared by four species. A transformation of main ornamental features (e.g. from punctate to smooth) can occur among closely related species suggesting short evolutionary timescales. Our overall results support the taxonomic value of carapace features, which should also receive greater attention in the taxonomy of extant species. The extensive variation in carapace shape and ornamentation is noteworthy and several species would probably have been assigned to different genera or families if these had been fossils, bearing implications for the systematics of fossil Spinicaudata.     

Multiple origins of a unique pollen feature: stellate pore ornamentation in Amaranthaceae

Stellate pore ornamentation is an unusual feature of angiosperm pollen, so far it is known in only ten genera of Amaranthaceae. The pollen grains of these plants have apertures with large hook‐shaped ektexinous bodies that are stellately arranged. Previous studies interpreted this character complex as a synapomorphy in consideration of its strong specialization. By reconstructing the evolution of stellate pore ornamentation based on phylogenetic trees of Amaranthaceae obtained by parsimony, likelihood, and Bayesian analysis of chloroplast trnK/matK DNA sequences, clear evidence is provided for several independent origins and reversals to less specialized aperture types. In addition to the gomphrenoid genus Pseudoplantago, stellate pore ornamentation evolved several times among achyranthoid genera, which have an African distribution. The most derived apertures, with 5 – 6 large protruding hooks, appear independently in Centemopsis on the one hand, and in Psilotrichum sericeum on the other. In an effort to break down the complex character syndrome of stellate pore ornamentation, we delimited a set of six pollen morphological characters that could be independently traced on the phylogeny. It appears that stellate pore ornamentation was independently derived from apertures with equally spread ektexinous bodies that became hook‐shaped, reduced in number, and symmetrically arranged. Likewise the symmetrically arranged, rectangular ektexinous bodies in Marcelliopsis represent an independent specialization. In view of this pattern of morphological changes, functional significance in the context of specialized insect pollination syndromes and positive selection for stellate pore ornamentation is hypothesized. Stellate pore ornamentation provides an example of a specialized pollen character complex with adaptive significance, and underlines the need for a dense taxon sampling for analyses of character evolution.     

Columellar muscle of neogastropods: muscle attachment and the function of columellar folds

Malacologists often assume that ornamentation on snail shells is functional, and therefore adaptive. I conducted the first comprehensive test of the widely accepted hypothesis that columellar folds, a type of internal ornamentation, enhance the performance of the columellar muscle, which attaches the snail to its shell. Careful dissections of live, non-relaxed specimens reveal that the physical attachment between the columellar muscle and the columella is not restricted to a small, circular patch located deep within the shell. Instead, the attachment is long and narrow, extending approximately a full whorl along the length of the columella. I developed a novel technique for preparing three-dimensional reconstructions from photographs documenting the dissections. These reconstructions were then used to measure four parameters that describe the muscle: (1) the surface area of the physical attachment between the muscle and columella, (2) the total contact area between the muscle and the columella, (3) the depth of attachment, and (4) the length of attachment. None of these parameters differed significantly between species with and without folds. In light of the biomechanics of muscular hydrostats, values of the first parameter indicate that columellar folds probably do not guide the columellar muscle as the animal moves in and out of its shell. Values of the other parameters indicate that columellar folds neither increase an animal's ability to maneuver its shell nor facilitate deeper withdrawal. These results, and the fact that folds have evolved convergently several times, might indicate that folds are an easily evolvable solution to many functional problems, none of which are currently understood.     

Comparative egg morphology of six Meccus species and Triatoma recurva (Stål, 1868) Hemiptera: Reduviidae

We investigated the morphology and morphometry of eggs from the colonies of the Entomology Laboratory at ENCB‐IPN belonging to six species of the genus Meccus and Triatoma recurva, using both light and scanning electron microscopy. Egg ornamentation is an important parameter to consider in the differentiation of species. Samples were observed under a scanning electron microscope. Measurements of fifty eggs per species included length, width, and opercular diameter, which were used for the morphometric analysis. The results showed that the seven species display a polygonal ornamentation dominated by hexagons; the operculum shows also a polygonal ornamental characteristic in each species. Morphometry provided valuable information for discriminating between closely related species of the genus Meccus and Triatoma recurva, a species akin to this group, thereby facilitating the complete discrimination of these species.     

山东水龙骨科植物孢粉学研究及其在分类上的意义

基于Flora of China采用国际石松类和蕨类植物新分类系统,其水龙骨科(Polypodiaceae)不仅涵盖了秦仁昌分类系统中的水龙骨科(Polypodiaceae) 20多属,而且还包括槲蕨科(Drynariaceae)、鹿角蕨科(Platyceriaceae)等独立的科。水龙骨科(Polypodiaceae)新分类系统种类较多、分类复杂,需要对其合理性和孢粉学在新分类系统中的意义进行新的探讨。该文首次以新分类系统为依据,采用扫描电镜对山东分布的水龙骨科(Polypodiaceae)植物孢子形态及其周壁纹饰亚显微结构进行了系统地观察。结果表明:该科孢子形态为圆肾形,左右对称,具周壁,其周壁纹饰在种内稳定,在亚科、属及种间区别显著。孢粉学研究结果,支持槲蕨亚科、鹿角蕨亚科分别列于水龙骨科下的两个亚科;基于山东假瘤蕨孢子周壁纹饰,并结合叶片形态特征,山东假瘤蕨(Phymatopteris shandongensis J.X.Li et C.Y.Wang)应为一个独立的新种,不宜并入金鸡脚假瘤蕨[Phymatopteris hastata (Thunb.) Pic.Serm.]。依据《国际植物命名法规》和Flora of China分类系统,山东假瘤蕨(Phymatopteris shandongensis J.X.Li et C.Y.Wang)新组合为山东假瘤蕨[Selliguea shandongensis (J.X.Li et C.Y.Wang) J.X.LiX.J.Li,comb.nov.]。该研究结果不仅首次为水龙骨科植物孢粉学积累了新资料,而且也为其新分类系统提供了孢粉学的科学依据,与DNA分类系统相吻合,证明其新分类系统的合理性和科学性,在构建水龙骨科及其亚科的自然分类系统中具有重要意义。