rbcL sequences support exclusion ofRetzia,Desfontainia, andNicodemia from theGentianales

The taxonomic positions ofRetzia, Desfontainia, andNicodemia have been much discussed, and all three genera have been included inLoganiaceae (Gentianales). We have made a cladistic analysis ofrbcL gene sequences to determine the relationships of these taxa toGentianales. Four newrbcL sequences are presented; i.e., ofRetzia, Desfontainia, Diervilla (Caprifoliaceae), andEuthystachys (Stilbaceae). Our results show thatRetzia, Desfontainia, andNicodemia are not closely related toLoganiaceae or theGentianales. Retzia is most closely related toEuthystachys and is better included inStilbaceae. The positions ofDesfontainia andNicodemia are not settled, butDesfontainia shows affinity for theDipsacales s.l. andNicodemia for theLamiales s.l.     

Phylogeny of theCyclanthaceae

The affinities ofCyclanthaceae are discussed, and it is concluded that the sister-group to this family is most probablyPandanaceae. A hypothesis of generic relationships inCyclanthaceae, based on cladistic methods, is presented. Bootstrap analysis and Bremer support (decay index) have been used to test the strength of individual clades, and the result is compared with previously made phylogenetic analyses. TheSphaeradenia group (Chorigyne, Stelestylis, Sphaeradenia, andLudovia) is supported as monophyletic and acceptably resolved, while theAsplundia group (remaining genera inCarludovicoideae) may be paraphyletic, with largely uncertain relationships. A formal recognition of these groups is therefore not justified. The probable character evolution inCarludovicoideae is discussed.     

P-type sieve-element plastids present in members of the tribesTriplareae andCoccolobeae (Polygonaceae) renew the links between thePolygonales and theCaryophyllales

Form-Pfs sieve-element plastids were found inTriplaris, Ruprechtia, andCoccoloba (Polygonaceae) while other genera of the family and those studied from the often associatedPlumbaginaceae contain S-type sieve-element plastids. The rareness of form-Pfs plastids among the angiosperms, their similarity to the peculiar form-P3fs plastids of theChenopodiineae, and the comparatively small plastid diameters measured for all forms present in theCaryophyllales, Polygonales, andPlumbaginales suggest close relationships between these taxa. The restriction inPolygonaceae of form-Pfs plastids to the closely allied tribesTriplareae andCoccolobeae is discussed with regard to both the intrafamilial and ordinal phylogeny, and also considering possible connections to the only magnoliidaean Pfs-taxonCanella. Dedicated to Univ.-Prof. DrF. Ehrendorfer on the occasion of his 70th birthday.     

Phylogenetic relationships of Scaphyglottis and related genera (Laeliinae: Orchidaceae) based on nrDNA ITS sequence data

Sequences of the nuclear ribosomal internal transcribed spacer regions 1 & 2 (nrDNA ITS) including the intervening 5.8S region were analyzed cladistically for 43 individuals of 35 species ofScaphyglottis s.l. plus two outgroup taxa. Low levels of sequence divergence do not allow estimation of relationships among most clades, but the analyses indicate that four segregate genera (Hexisea Lindl.,Reichenbachanthus Barb. Rodr.,Hexadesmia Brogn., andPlatyglottis coriacea L.O. Williams) are embedded within a broad paraphyleticScaphyglottis. This broadly definedScaphyglottis sensu Dressler is characterized within Laeliinae by the usual presence of superposed growth habit and the presence of a column foot. In order to accommodate species formerly placed inPlatyglottis andReichenbachanthus, three new combinations are made inScaphyglottis:Scaphyglottis brasiliensis (Schltr.) Dressler,S. coriacea (L. O. Williams) Dressier, andS. emarginata (Garay) Dressler.     

Phylogenetic relationships of epacrids and vaccinioids (Ericaceae s. l.) based onmatK sequence data

Cladistic relationships of epacrids and vaccinioids (Ericaceae) are investigated using nucleotide sequence data from the chloroplast encodedmatK gene. Sequences of 56 taxa were aligned and analyzed using parsimony methods. Results show thatVaccinioideae as currently recognized are not monophyletic. The epacrids are sister to a clade that includes theLyonia group, theGaultheria group, and theVaccinieae. Arbutus andPyrola branch early inEricaceae, before the rhododendroid group.Enkianthus is sister to the remainingEricaceae (includingEpacridaceae).Vaccinieae are strongly supported as monophyletic, butVaccinium andAgapetes are polyphyletic.     

Some Entolomataceae (Agaricales) from Costa Rica

A preliminary account of Entolomataceae from Costa Rica is presented. Three new taxa are described, two inClitopilus, one inRhodocybe, and three new combinations are proposed inInocephalus. Five taxa ofClitopilus are reported for the first time from Costa Rica and a key to species is provided. Four species ofRhodocybe are discussed and a key to the six species known from Costa Rica is also provided.Alboleptonia earlei, Inocephalus murraii, Inocephalus quadratum, Rhodocybe incarnata andRhodocybe pseudonitellina are now known to occur in Costa Rica.     

Karyosystematic studies on threeCrepis species (Asteraceae) endemic to Greece

This work examines the cytogeographical distribution, the morphological characters, and the karyotypes of threeCrepis species endemic to Greece (C. sibthorpiana, C. incana, andC. heldreichiana). C. sibthorpiana is diploid (2n = 2x = 8),C. incana is diploid (2n = 2x = 8) and tetraploid (2n = 4x = 16, 17), andC. heldreichiana is always dekaploid (2n = 10x = 40). The Giemsa positive bands, usually pairs of dots, are mainly centromeric inC. incana, while they are terminal inC. sibthorpiana (on the short arm of all chromosomes) and inC. heldreichiana (on both arms of all chromosomes). Intercalary C-bands are scarce and usually variable within karyotypes, individuals, and species. The most variable karyotype both in Feulgen and Giemsa preparations is that ofC. incana, in which also supernumerary chromosomes were observed, which are polysomic to standard set members. On the basis of morphological and karyological data the evolutionary relationships among the threeCrepis taxa are discussed.     

Floral development and systematics ofCistaceae

The floral development of representatives of six genera ofCistaceae has been studied. Calyx development involves the formation of a ring primordium in several taxa. Androecium development in species with intermediate or higher stamen numbers starts with the formation of a ring meristem on which the stamens are initiated in a centrifugal direction. In many taxa five alternipetalous leading stamen primordia can be observed. In the apetalous (cleistogamous) flowers ofTuberaria inconspicua androecium development appears to be unordered; this is probably due to the lack of petals. InLechea intermedia (also cleistogamous) the corolla is trimerous and three complex stamen primordia are produced, which give rise either to one or three stamens. Relationships withinCistaceae are discussed. Floral development inCistaceae is compared with that in otherMalvanae. Among the eight families ofMalvanae from which information on floral development is availableCochlospermaceae andBixaceae exhibit the greatest similarities toCistaceae. InCistaceae the leading stamen primordia are alternipetalous. InBixa the same condition seems to be present. InMalvales s. str. mostTiliaceae also show earliest stamen initiation in alternipetalous sectors, whereas the stamens of the innermost alternipetalous position are retarded early or even suppressed inSterculiaceae, Bombacaceae, andMalvaceae. WithinMalvales s. str. the diversity of androecial developmental patterns seems to decrease inBombacaceae andMalvaceae due to increasing synorganization in the mature androecium. The derivation of polyandry inMalvanae from diplo- or obdiplostemony is discussed by comparison with the sister clades ofMalvanae as shown in recentrbcL studies (i.e.Sapindales, Rutales, the glucosinolate producing clade, andMyrtales).     

Electrophoretic seed albumin patterns and species relationships inVicia sect.Faba (Fabaceae)

Electrophoretic analysis of seed albumins (PAGE) covered 173 accessions representing nine species ofVicia sect.Faba. The number of albumin bands recorded in particular species varied from three inV. eristaloides to 23 inV. faba; in total, 38 bands were distinguished in the investigated material. The examined species, exceptV. eristalioides, showed intraspecific variation with respect to the number and relative staining intensity of albumin bands; individual variation was especially marked inV. faba and inV. narbonensis. Hierarchical clustering of the investigated taxa was based onBhattacharyya distances calculated from the electrophoretic data. The taxa grouped in three main clusters.Vicia faba and the rather remotely relatedV. kalakhensis formed one cluster. The second cluster was composed ofV. narbonensis distantly related toV. hyaeniscyamus. The third cluster comprised three subgroups: 1.V. johannis, V. galilaea andV. serratifolia, 2.V. eristalioides, and 3.V. bithynica. The obtained results are discussed with reference to taxonomic relationships inVicia sect.Faba.     

Identity of North African endemic bryophytes, 1

Abstract

Ten taxa of Pottiaceae considered endemic to North Africa have been studied and typified. Nine are newly synonymized with other well-known taxa. A new combination is proposed: Weissia condensa var. armata (Thér. & Trab.) M.J.Cano, Ros & J.Guerra. The distribution of Acaulon triquetrum var. desertorum (Besch.) Jelenc and Weissia condensa var. armata are respectively extended to Europe and the African continent. Barbula muralis var. obovata Schimp. is a misprint in the publication of the taxon. Two nomina nuda, Tortula cuneifolia var. caulescens Jelenc and Tortula cuneifolia var. pilifera Jelenc have no taxonomic value.     

Hybridisation,genomic constitution and generic delimitation inElymus s. l. (Poaceae: Triticeae)

Intergeneric crosses were made between representatives of the genomically-defined generaElymus, Agropyron, Elytrigia, Pseudoroegneria, andThinopyrum. The genomic constitution ofElytrigia repens, the type species ofElytrigia, is shown to be SSH, a genomic combination otherwise found only inElymus. The S genome ofPseudoroegneria has almost always a dominant influence on the morphology of the taxa of which it is a component.Wang (1989) showed that the J genome inThinopyrum and the S genome have considerable homoeology, with a mean c-value of 0.35 in diploid SJ hybrids. A genetic coherence from S to SJ^e, J^e, J^eJ^b, and J^b can be expected, agreeing with the continuous morphologic variation pattern observed. Because of the absence of morphological discontinuities between the taxa,Pseudoroegneria (S),Elymus (SH, SY, sometimes with additional genomes),Elytrigia (SSH, SSHX), andThinopyrum (SJ, SJJ, J) are best treated as a single genus,Elymus, following the generic concept ofMelderis in Flora Europaea and Flora of Turkey. The basic genomic constituents ofElymus will then be the S and/or J genomes.Agropyron, with diploids, tetraploids, and hexaploids based on the P genome is morphologically distinct from other genera inTriticeae. In a few species ofElymus andPseudoroegneria, a P genome is an additional constituent. In these cases the P genome has a negligible morphological influence. Therefore, it seems reasonable to maintainAgropyron as a separate genus.     

Phylogenetic analysis of the generaCladonia andCladina (Cladoniaceae, lichenizedAscomycota)

A cladistic analysis of 44 species of the generaCladina andCladonia is presented.Pycnothelia papillaria, Cladia aggregata andC. retipora were used as outgroup taxa. The consensus of all equally parsimonious trees suggests a common ancestral origin for species inCladonia andCladina while the generaPycnothelia andCladia cluster outside this group. The results do not support distinction ofCladina at genus level although it is distinguished as a monophyletic group but within the genusCladonia. The current sectional division ofCladonia is not supported. SectionsCocciferae andHelopodium are best supported but even these groups as currently delimited seem to in include elements that are more closely related to species of the other sections, e.g.C. carneola of sect.Cocciferae andC. pityrophylla of sect.Helopodium. Sections such asUnciales andPerviae seem to be artificial assemblages. In general the evolutionary scheme of the genus seems to be more complicated than revealed by the current sectional division.     

Pollen dimorphism in threeTeucrium species (Lamiaceae)

In the Iberian Peninsula and Balearic Islands pollen dimorphism was found inTeucrium fruticans L.,T. pseudochamaepitys L., andT. rotundifolium Schreber. Generally, this dimorphism shows two sizes of pollen grains, the smaller more or less collapsed. The percentage of pollen viability is calculated. Differences in size between viable and nonviable pollen grains are similar, being about 40% inT. fruticans andT. pseudochamaepitys, and about 26% inT. rotundifolium. With regard to pollen viability, the percentage of male sterility is higher inT. fruticans, in which from male sterile (ms) plants, 100% nonviable pollen was obtained from every flower observed. InT. pseudochamaepitys andT. rotundifolium with rare exceptions, the percentage of nonviable pollen does not seem to be significant.     

Utility of rDNA ITS sequences in the systematics ofTeucrium sectionPolium (Lamiaceae)

A phylogenetic study based on sequence data from the complete internal transcribed spacer region (ITS) of nuclear ribosomal DNA for sect.Polium of the genusTeucrium shows both intersectional congruence and intrasectional incongruence with traditional morphological classifications. We attribute this incongruence largely to problems related to the polyploid complex studied. SectionPolium includes many poorly differentiated taxa of probable recent origin through hybridization followed by polyploidization. Both on the basis of parsimony and distance (Neighbor-Joining method) analyses,T. dunense andT. thymifolium are the species that diverge most from the sampled taxa. However, unlikeT. thymifolium, the morphology ofT. dunense is not much differentiated in relation to the other taxa. Both species are, nonetheless, the only sampled species to occupy isolated, exclusive environments and which may have undergone rapid evolution by a bottleneck effect.Teucrium dunense is found only on dunes along the Spanish and French coasts andT. thymifolium, a chasmophytic species with limited endemism, is found solely on limestone and dolomite cliffs in the low mountains in south-eastern Spain. A hypothesis is presented to explain the discrepancy between the observed comparatively large amount of ITS sequence divergence and the low morphological differentiation inT. dunense.     

High Mitochondrial DNA Diversity with Little Structure Within and Among Leaf Monkey Populations (Trachypithecus cristatus and Trachypithecus auratus)

We used analyses of mitochondrial DNA restriction site polymorphisms to estimate population genetic structure and phylogenetic relationships among 42 individuals from two Asian leaf monkey species (Trachypithecus auratus and T. cristatus) and to compare them to the geographically proximate species, Presbytis comata. We amplified a 2300-base pair fragment spanning the mitochondrial NADH 3 and NADH 4 genes, including their tRNA flanking subunits, glycine and leucine, and digested it with a battery of 22 restriction endonucleases, yielding 21 unique multienzyme haplotypes and 60 variable restriction sites. Presbytis comata is clearly divergent from both Trachypithecus species. Within the Javan T. auratus, our analysis does not support the distinction of two subspecies currently recognized on the basis of morphological features (Weitzel and Groves, 1985). T. auratus and T. cristatus are not internally monophyletic with respect to each other in our phylogenetic analysis. These results indicate either a recent speciation event with the retention of ancestral polymorphisms or that the two taxa are not separate species. Therefore with respect to conserving genetic diversity within the leaf monkey, we would have to consider T. auratus and T. cristatus as essentially one large polymorphic, conservation unit. However, within that conservation unit, T. auratus of Java represent a separate management unit from T. auratus/T. cristatus of Sumatra and Peninsular Malaysia.     

Phylogeny ofRubiaceae-Rubieae inferred from the sequence of a cpDNA intergene region

A phylogenetic analysis of 25 species, representing eight genera of theRubieae tribe (Rubiaceae), has been made using the DNA sequence of the chloroplastatp B-rbc L intergene region. Six tropical genera from other tribes ofRubiaceae have been used as outgroups. Whatever the method of analysis (distance, parsimony or maximum likelihood), five groups are clearly separated and described as informal clades. Their relative relationships are not clearly resolved by the parsimony analysis, resulting in eight equally parsimonious trees, 327 steps long, with a consistency index (CI) of 0.749 (excluding uninformative sites). TheRubieae tribe appears monophyletic from the data available. Some new and partly unexpected phylogenetic relationships are suggested. The genusRubia forms a separate clade and appears to be the relatively advanced sister group of the remaining taxa. TheSherardia clade also includes the generaCrucianella andPhuopsis. Galium sect.Aparinoides appears closely attached to theAsperula sect.Glabella clade. The remaining taxa ofGalium are paraphyletic:Galium sect.Platygalium (in theCruciata clade) is linked to the advanced generaCruciata andValantia; the more apomorphic groups ofGalium form theGalium sect.Galium clade, including the perennial sectionsGalium, Leiogalium, andLeptogalium as well as the annual (and possibly polyphyletic) sect.Kolgyda.     

Implications of morphological phylogenetics for the placement of the generaAdenocaulon andEriachaenium (Asteraceae)

Adenocaulon andEriachaenium are two problematic genera because their tribal and subfamilial placement in Asteraceae is uncertain. Previous cladistic analyses based on molecular data undertaken to analyze the relationships within Asteraceae, placeAdenocaulon in the tribe Mutisieae (Cichorioideae). This paper investigates cladistic relationships amongAdenocaulon andEriachaenium and tribes of subfamilies Cichorioideae and Asteroideae using morphological data. Thirty-eight characters were scored across 52 genera selected as exemplar taxa to represent the current classification system. In the analysis (one tree, length = 86, c.i. = 0.55, r.i. = 0.64)Adenocaulon andEriachaenium are sister taxa and appear as an isolated clade nested in Cichorioideae. A new, tentative position among the tribes of the paraphyletic Cichorioideae is proposed for these two isolated genera.     

Variation in wild populations of annual beet (Beta,Chenopodiaceae)

Among the morphological variation in wild annual populations of sect.Beta only tepal characters show a geographic pattern, and hence can be used to distinguish different taxa. Two morphological types correspond with taxa already described:B. macrocarpa (incl.B. bourgaei) has a Macaronesian and W. to E. Mediterranean andB. adanensis an E. Mediterranean distribution area. A third type in the Aegean region is not well known yet and possibly has to be included inB. macrocarpa. Both diploid and tetraploid (x = 9) cytotypes are found withinB. macrocarpa, the latter exclusively on the Canary Islands.     

Studies on the embryology and gynoecium structures inDrimys winteri (Winteraceae) and someAnnonaceae

Drimys winteri (Winteraceae) and 11 species ofAnnonaceae, namelyAnnona montana, Artabotrys hexapetalus, Bocagea sp.,Papualthia sp.,Polyalthia nitidissima, Tetrameranthus umbellatus, T. duckei, Uvaria sp.,Xylopia malayana, X. aromatica, andX. emarginata, were investigated embryologically with special reference to development of ovule and embryo sac. The ovules are anatropous, crassinucellate and in most taxa bitegmic. The inner integument is of epidermal origin. TheAnnonaceae investigated have a multi-layered, later vascularized outer integument with most probably subepidermal initiation. In contrast,Drimys winteri has a three-layered, non-vascularized outer integument of epidermal origin. The annonaceous genusTetrameranthus (T. umbellatus andT. duckei) possesses a middle integument between the inner and the outer one, stated here for the first time in a neotropic representative ofAnnonaceae. Within the angiosperms this feature occurs inAnnonaceae only. The embryological characters are rather homogeneous. Differences between the species investigated are found in, e.g. the number of cell layers in the inner integument, which is commonly two inAnnonaceae as compared to three inDrimys winteri, the presence or absence of a hypostase, the number of layers in the nucellar epidermis, great differences in size of ovules, and the species-specific pattern of tannin deposition in the ovules. In the species so far investigated the embryo sacs develop according to thePolygonum-type. InXylopia malayana andBocagea sp. in addition the carpels were investigated. They are conduplicate. InXylopia malayana the free carpels are united by an extragynoecial compitum, inBocagea sp. each stigma produces its isolated mucilage cap. The results obtained from the investigated taxa are discussed and compared with published data on embryology and gynoecium structure in other annonaceous and winteraceous taxa.     

Morphological analysis ofAlouatta seniculus species group (Primates,Cebidae). A comparison with biochemical and karyological data

Fifteen cranial measurements were taken from wild caught specimens ofAlouatta seniculus seniculus, A. s. stramineus andA. s. macconnelli. A morphological analysis showed sex dimorphism in these three groups. A multivariate analysis discriminated among these taxa; males being more clearly discriminated than females. Our data showed that these taxa can be separated on the basis of quantitative cranial traits, biogeographic distribution, karyological differences, and biochemical characters. We therefore propose a new taxonomic arrangement, changing their taxonomic status to the species level (Alouatta seniculus, A. macconnelli, andA. stramineus).