Biometrische Untersuchungen an Populationen vonOphrys cornuta,O. holosericea und ihren Hybriden (Orchidaceae)

Population variability ofOphrys holosericea (Burm. f.)Greut. subsp.holosericea (=O. fuciflora Crantz subsp.fuciflora) from near Vienna (Austria), and of subsp.maxima (Fleischm.)Greut. andO. cornuta Steven with intermediates from the Dalmatian island Hvar (Yugoslavia) was analysed and illustrated by scatter diagrams. A hybrid origin of these intermediates is suggested. Aspects of hybridization betweenO. holosericea agg. andO. scolopax agg. are discussed.

     

Populationsstatistische Analysen bestätigen die Heterogeneität vonOphrys „arachnitiformis“ (Orchidaceae)

Statistical population analyses have been carried out on so-calledOphrys arachnitiformis, partly sympatricO. sphecodes s. l. as well as on other species ofOphrys (O. holosericea, O. exaltata, O. apulica, O. cornuta, O. biscutella, O. bertolonii, O. bertoloniiformis.). Altogether 63 populations with 521 individuals were studied in regard to 54 morphological parameters. — It can be shown thatO. arachnitiformis is an extremely heterogeneous assemblage which includes taxa of most diverse origin, different degrees of stability and nomenclatorial rank. Only common trait is their very diverse relationships withO. specodes s.l.—The basionymO. arachnitiformis Gren. & Phil. refers to a colour variant ofO. sphecodes Mill. subsp.sphecodes only, and is not applicable to other segments of the assemblage.A late flowering, morphologically and biologically well isolated group of populations from southern France without any perceptible hybrid influence is established as a new species:Ophrys splendida Gölz & Reinhard. Populations from north and central Italy with distinct characters but hybrid influences fromO. holosericea/O. exaltata are calledO. tyrrhena Gölz & Reinhard.O. morisii, an endemic to Sardinia, represents an independent species of still obscure evolution. The hybrid interpretation ofO. catalaunica is statistically corroborated, but somewhat modified. For the remaining segments ofO. arachnitiformis investigated (from Castilia, Dalmatia, and the Monte Gargano) no formal taxonomic proposals are made, but some suggestions about their genesis are possible.

     

Neue Beobachtungen zur Bestäubung vonOphrys (Orchidaceae) in Südspanien,mit besonderer Berücksichtigung des FormenkreisesOphrys fusca agg.

An investigation of pseudocopulation behaviour in species ofOphrys from southern Spain confirms the close relationship betweenCampsoscolia ciliata (Scoliidae) andOphrys speculum, and betweenEucera nigrilabris (Apoidea) andOphrys tenthredinifera. It could be demonstrated thatEucera barbiventris is the pollinator ofOphrys scolopax subsp.scolopax, whereas other species ofEucera andTetralonia which are active at the same time show no interest at all for the flowers of this species. Special attention was paid to the forms of the taxonomically confusedOphrys fusca group:O. fusca s. str.,O. iricolor, O. omegaifera andO. atlantica. WhileO. fusca s. str. is widespread, small-flowered and has late anthesis,O. iricolor has very large flowers and early anthesis. Each of the four members ofO. fusca agg. in S. Spain is pollinated by a different bee, and selective experiments show that each of these four species of bees is specifically attracted only to one of theOphrys species:Andrena flavipes is the pollinator ofO. fusca s. str.,Colletes cunicularius infuscatus ofO. iricolor, Anthophora atroalba ofO. omegaifera, andChalicodoma parietina ofO. atlantica. These four pollinators belong to 4 different bee families (Andrenidae, Colletidae, Anthophoridae, Megachilidae). As this type of pollination represents a very effective pregamic isolation mechanism, these four taxa ofOphrys fusca agg., at least in southern Spain, behave and should be regarded as genuine species.

     

Gene variation and gene flow inOrchis morio (Orchidaceae) from Italy

Data are presented on genetic variation at 27 enzyme loci of the Green-Winged orchid,Orchis morio, in 18 population samples from Italy. The existence in Italy of two subspecies, i.e. subspp.morio andpicta, is not supported by allozyme data. No genetic heterogeneity was found betweenmorio-like andpicta-like samples and specimens. Moreover, morphological transition between the two forms was observed in different Italian populations. The parameters of genetic variability estimated forO. morio populations are consistent with those found among monocotyledon plants, and among those outcrossing, animal-pollinated and with wind-dispersed seeds. Genetic diversity of ItalianO. morio is mostly within populations. Correspondingly, low values of interpopulational genetic distance were found. This appears to be due to high levels of gene flow, which were estimated with different methods. The lack ofO. longicornu from Italian samples, as well as of any hybrid withO. morio (F₁, backcrossed or recombinant individuals) is demonstrated on the basis of genetic data. It is concluded that recurrent reports ofO. longicornu from Italy are due to confusion withO. morio or with otherOrchis species.     

Ophrys helenae (Orchidaceae), eine lange verkannte Sippe der Balkanhalbinsel

Population analyses, scatter diagrams and field observations demonstrate thatOphrys helenae is morphologically (and ecologically) well separated from the partly sympatricO. mammosa andO. ferrumequinum in N. Greece. Hybrids with the distantly relatedO. mammosa are very rare.

     

Reproductive isolation in the Aegean Ophrys omegaifera complex (Orchidaceae)

The orchid genus Ophrys operates a system of sexual deception by which high specificity of pollination is attained. Reproductive isolation in Ophrys mainly rests upon prezygotic isolation mechanisms. The level of genetic separateness of Ophrys taxa with different pollinators is therefore likely determined by the fidelity of pollinators. The present study employs genetic fingerprinting to investigate this in the east Aegean Ophrys omegaifera s.l. complex, also including O. dryis, a west Mediterranean species of this complex. Ophrys fleischmannii, O. basilissa, and the west Mediterranean O. dyris, are found to be well-separated genetic entities whereas O. omegaifera s.str. and the putative hybrid taxon, O. sitiaca, are found to be genetically inseparable across their entire range of co-occurrence. This suggests that specific pollinators have high enough fidelity to act as effective isolating factors in east Aegean O. omegaifera s.l. as a whole, but that the situation in the species pair of O. sitiaca and O. omegaifera is likely to be more complex.     

Pollination ofOphrys (Orchidaceae) in Cyprus

In the southern part of Cyprus the pollinator —Ophrys (Orchidaceae) relationships and its specifity have been investigated from the end of February until the middle of March 1986. 12Ophrys spp. were found. To date, only a single pollinator reference has been reported from this island. We found the following pollinators:Melecta tuberculata (Ophrys kotschyi),Eucera dimidiata (Ophrys flavomarginata),Eucera gaullei (Ophrys umbilicata),Eucera paulusi (Ophrys bornmuelleri),Anthophora erschowi (Ophrys elegans),Andrena torda (Ophrys sicula =O. lutea subsp.minor),Andrena cinereophila (Ophrys fusca, small-flowered),Andrena flavipes (Ophrys israelitica),Andrena morio (Ophrys iricolor andOphrys transhyrcana),Andrena bimaculata (Ophrys sphegodes aggr., probably formerly confused withO. transhyrcana). Most interestingly, it could be verified thatO. flavomarginata/O. umbilicata, O. bornmuelleri/O. levantina andO. transhyrcana/O. sphegodes aggr. (possiblyO. sintenisii) are different biospecies. This is a result of genetic isolation due to varying pollinators, and of differences in flower morphology.     

Labellum micromorphology in theOphrys bertolonii agg. and some related taxa (Orchidaceae)

Micromorphology and distribution of trichomes in six taxa of theOphrys bertolonii aggregate (O. bertolonii, O. bertoloniiformis, O. benacensis, O. aurelia, O. drumana, O. catalaunica) are described. A fairly good homogeneity between all the taxa, with a moderate degree of variability even within the same taxon, has been observed. Comparison of these characters with those ofO. promontorii andO. garganica corroborates the inclusion of the former species into theO. sphegodes aggregate. Trichome analysis of a hybrid is exemplified byO. ×chiesesica (O. drumana ×O. fuciflora).     

Polyploidy and aneuploidy inOphrys,Orchis, andAnacamptis (Orchidaceae)

Studies on chromosome numbers and karyotypes in Orchid taxa from Apulia (Italy) revealed triploid complements inOphrys tenthredinifera andOrchis italica. InO. tenthredinifera there is no significant difference between the diploid and the triploid karyotypes. The tetraploid cytotype ofAnacamptis pyramidalis forms 36 bivalents during metaphase I in embryo sac mother cells. Aneuploidy was noticed inOphrys bertolonii ×O. tarentina with chromosome numbers n = 19 and 2n = 38. There were diploid (2n = 2x = 36), tetraploid (2n = 4x = 72), hexaploid (2n = 6x = 108) and octoploid (2n = 8x = 144) cells in the ovary wall of the diploid hybridOphrys apulica ×O. bombyliflora. Evolutionary trends inOphrys andOrchis chromosomes are discussed.     

Molecular (RAPD) analysis on Italian taxa of theOphrys bertolonii aggregate (Orchidaceae)

The RAPD technique has been used for analyzing the genetic material of seven Italian populations of theOphrys bertolonii aggregate (Orchidaceae). For comparison the related speciesO. sphegodes andO. fuciflora have also been analyzed. A high genetic variability has been observed within the same populations, and on the whole onlyO. bertolonii can be safely differentiated from the other taxa of the aggregate, which are undistinguishable with this method.     

Zwergstrauchige Arten der FlechtengattungCaloplaca

The frutescent species of the lichen genusCaloplaca are usually united in sect.Thamnoma, but they do not form a natural group. They are derived from different species groups within sect.Gasparrinia from different parts of the world, presumable from species having scleroplectenchymes in cortex and medulla. The algal cells are concentrated between the scleroplectenchymatic strands in large and dense groups, from where medullary plectenchyme extends to the cortex and forms characteristic pseudocyphellae there.—Most of the species seem to be ornithocoprophilous; they grow on rocks along marine coasts where much fog is induced by cold currents.—Caloplaca cribrosa is endemic in Tasmania and New Zealand,C. regalis and the doubtfulC. ambitiosa belong to the antarctic element.C. fragillima from central Chile seems to be propagated by thallus fragments.C. coralloides andC. thamnodes are endemic to California and Baja California respectively.C. cladodes from the Rocky Mountains deviates in many characteristics from the other species i.a. by it different ontogenetic development, reduced spore septum, and cementing amyloid polysaccharides within the scleroplectenchymatic strands. The African species are characterized by their distinctly dorsiventral lobes and usually possess oil cells in some of the paraphyses.Caloplaca bonae-spei, C. fragillima andC. thamnodes are new to science.

     

Crossing and selfing experiments in theOrnithogalum umbellatum/augustifolium complex

The fertility relationships ofOrnithogalum umbellatum, O. angustifolium and 9 related taxa were studied.O. umbellatum andO. angustifolium, which are not interfertile, are both recognized as a comparium. Pollination of five species belonging to theO. umbellatum/angustifolium polyploid complex, i.e.O. umbellatum, O. angustifolium, O. baeticum, O. monticolum andO. algeriense, resulted in a moderate seed set. However, the seeds obtained after these interspecific crosses did not germinate. Six related species, i.e.O. exscapum, O. nivale, O. comosum, O. collinum, O. gussonei andO. woronowii showed a very low seed set after being crossed. One exception occurs:O. nivale ×O. exscapum. The seed set was also low after the complex species had been crossed with some related species, but there was one exception: both reciprocal crosses betweenO. monticolum andO. woronowii yielded some viable seeds. No other interspecific crosses gave rise to viable seeds. Bulbil-producing species have a lower seed set after self-pollination than species which do not produce bulbils.Biosystematic Studies on theOrnithogalum umbellatum/angustifolium Complex IV.—Previous contributions: (I–III):Raamsdonk (1982, 1985a, 1985b).     

Neue Taxa vonTillandsia subgenusDiaphoranthema (Bromeliaceae) aus Bolivien und Argentinien

5 new taxa are described and illustrated; their position within the subgenus is discussed:T. hirta W. Till & L. Hromadnik,T. cotagaitensis L. Hromadnik,T. caliginosa W. Till, andT. gilliesii Baker subsp.polysticha W. Till & L. Hromadnik are members of a group includingT. myosura Grisebach exBaker,T. mandonii E. Morren exMez in DC.,T. retorta Grisebach exBaker em.Grisebach andT. andicola Gillies exBaker. T. brealitoensis L. Hromadnik is related toT. angulosa Mez in DC. but distinct and possibly of hybrid origin.

     

RAPD characterisation of two neotropical hybrid legumes

An investigation of randomly amplified polymorphic DNA (RAPD) and polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) marker distribution was made for two well-characterised hybrids and their parents,Leucaena leucocephala andL. esculenta andParkinsonia aculeata andCercidium praecox. Three chloroplast DNA (cpDNA) markers identified the maternal parent of eachL. leucocephala ×L. esculenta hybrid. Fifteen species-diagnostic RAPD markers (invariant in one taxon and absent from the other) were always present in theLeucaena hybrid and assumed to be of nuclear origin, whilst three RAPD markers showed expression patterns identical to the cpDNA markers and were assumed to be of organellar origin. No RAPD or PCR-RFLP taxon-diagnostic markers were discovered for eitherP. aculeata orC. praecox. However, 21 RAPD markers were species-specific (polymorphic within one taxon but absent from the other) and Southern analysis indicated that none of the markers were of organellar origin. Only 67% additivity of markers specific toP. aculeata andC. praecox was demonstrated in the hybrids between these two species, whilst inLeucaena 97% additivity was demonstrated. Differences between the two hybridising situations were related to the behaviour of the molecular markers and the biology of the species.     

Production and cytogenetic analysis of intergeneric hybrids betweenElymus anthosachnoides andPsathyrostachys huashanica (Poaceae: Triticeae)

Intergeneric hybridization betweenElymus anthosachnoides (2n = 28,SSYY) andPsathyrostachys huashanica (2n = 14,NN) was performed. Three hybrid plants, obtained via embryo rescue, were intermediate between the parents in morphology and developed vigorously, but were completely sterile. The mean chromosome configuration was 19.48 I + 0.76 II per cell in the hybrids at meiotic metaphase I. This result indicates thatE. anthosachnoides andP. huashanica are distantly related and that there is little or no homoeology betweenN (P. huashanica) andS orY (E. anthosachnoides) genomes.     

Interspecific hybridization between the cultivated potato Solanum tuberosum subspecies tuberosum L. and the wild species S. circaeifolium subsp. circaeifolium Bitter exhibiting resistance to Phytophthora infestans (Mont.) de Bary and Globodera pallida (Stone) Behrens

Summary Somatic fusions between the cultivated potato Solanum tuberosum and the wild species S. circaeifolium subsp. circaeifolium Bitter were produced in order to incorporate desirable traits into the potato gene pool. Selection of the putative hybrids was based on a difference in callus morphology between the hybrids and their parents, with the hybrids showing typical purple-colored cells in otherwise green calli. In all, 17 individual calli regenerated to plants. Of the nine plants that could be transferred to the greenhouse, eight showed a hybrid and one a parental morphology. Restriction fragment length polymorphism (RFLP) analysis confirmed the hybrid character in the former group. Chloroplast counts in stomatal guard cells and flow cytometric determination of nuclear DNA content showed that four hybrid plants were tetraploid (4x), one was mixoploid (5x–8x), and the others were polyploid (6x; 8x). Three out of four tetraploid hybrids were found to be fully resistant to Phytophthora infestans, and all four hybrids were resistant to Globodera pallida pathotypes Pa2 and Pa3. It was further observed that the type and amount of steroidal glycoalkaloids varied among the tubers of the parents and the hybrids. Using the hybrids as female parents in crosses with S. tuberosum, viable seeds could be obtained. This demonstrates the potential of these hybrids in practical plant breeding.     

Isozyme evidence for the parentage and multiple origins ofPotamogeton ×suecicus (P. pectinatus ×P. filiformis,Potamogetonaceae)

Evidence from isozyme analyses indicates thatPotamogeton ×suecicus is the hybrid betweenP. pectinatus andP. filiformis. The hybrid appears to have arisen on several occasions. The isozyme profiles of this hybrid from the Rivers Wharfe and Ure in Yorkshire, south of the present limit of distribution ofP. filiformis, suggest that each population is a single clone; these clones may be relics from the Weichselian glacial period. Populations of the putative hybrid from the Rivers Tweed and Till are notP. ×suecicus but probably haveP. vaginatus andP. pectinatus as parents. If so, this is a remarkable example of a pondweed hybrid persisting vegetatively in an area outside the distributional range of one of its parents.     

Drei neue Arten desFestuca ovina-Formenkreises (Poaceae) aus dem Osten Österreichs

WithinFestuca ovina agg. two hexaploid taxa:F. brevipila andF. carnuntina, and one tetraploid:F. eggleri, are described as new species.F. pseudovina Hack. exWiesb. is better treated asF. valesiaca Schleich. exGaudin subsp.parviflora (Hack.), comb. nova.

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Drei neue Arten desFestuca ovina-Formenkreises (Poaceae) aus dem Osten Österreichs

     

The association between pollen size and Renner complex in Oenothera villaricae and O. picensis ssp. picensis and their hybrids: evidence for preanthesis pollen competition

In both Oenothera villaricae Dietrich and O. picensis ssp. picensis, chromosomes are transmitted as two Renner complexes. Reciprocal combinations of the Renner complexes produce eight different F₁ hybrids, but only seven are viable. Each species, and each F₁ hybrid, produces three sizes of pollen, approximately 50% small sterile grains, 15% medium-sized viable grains and 35% large viable grains. Medium- and large-sized grains were separated manually and subjected to random amplified polymorphic DNAs (RAPDs) analysis. A pattern of RAPD amplifications was obtained which indicates that, for each species and F₁ hybrid, one specific Renner complex characterizes the medium- and another the large-sized viable pollen. The results indicate that pollen size is determined in part by the pollen genotype and in part by the genotype of the other pollen grains developing within the same anther.     

Species novae generisEremostachys (Labiatae)

Five new species are described:Eremostachys codonocalyx from NE. Iran (related toE. subspicata),E. stenocalycina from NE. Afghanistan (related toE. alberti andE. hissarica),E. salangensis from NE. Afghanistan (close toE. bamianica),E. freitagii from E. Afghanistan (similar but not closely related toE. vulnerans), andE. andersii from E. Afghanistan (similar but not closely related toE. vulnerans), andE. andersii from NE. Afghanistan of isolated systematic position.

Florae Iranicae praecursores 5–9.