Morphological variation inLolium (Poaceae) as a measure of species relationships

Analysis of morphological and phenological data for determining the genetic variation within sevenLolium species led to the recognition of two groups within this genus. One group, containing the two inbreeding speciesL. temulentum andL. persicum, was clearly distinct from all other species. Strong morphological and phenological intergradation was found between both species. The cross-breeding species,L. perenne, L. rigidum, andL. multiflorum, formed another group. Little differentiation was found between these species, though they were distinct. Two inbreeding species,L. loliaceum andL. remotum, were clearly distinct from each other and the two groups.L. loliaceum had an isolated position and was most related toL. rigidum. L. remotum had an intermediate position between the cross-breeding and inbreeding species, and was almost equally distant from all three cross-breeding species.Genetic variation inLolium spp. I.     

Species relationships inFestuca (sect.Ovinae,Poaceae)

Morphological and seed protein studies of selected species ofFestuca were performed to elucidate the relationships between species of sect.Ovinae and their affinity with other fescues.Festuca rubra andF. heterophylla (extravaginale group) had higher phenetic affinity with taxa of sect.Scariosae, Montanae andBovinae than with members of their own section. 5 species of the intravaginale group (F. ovina, F. capillata, F. valesiaca, F. sulcata, F. trachyphylla) were fairly uniform and well defined by protein and morphological data. This supports current systematic thinking and does not agree withHackel's (1882) classification of this group.     

Chromosome banding patterns in Lettuce species (Lactuca sect.Lactuca,Compositae)

Chromosome banding patterns obtained with C- and N- banding, and AgNO₃ staining were studied in somatic metaphase complements of fourLactuca species.L. sativa andL. serriola have almost identical chromosome morphology, andL. saligna differs only slightly from them, butL. virosa is quite distinct from the other species. A gross comparison of the banded karyotypes suggests a closer relationship ofL. saligna toL. sativa/serriola than toL. virosa. Our data agree with the results of previous crossing experiments in these species but conflict partly with recent RFLP data which indicate a closer phenetic relationship ofL. saligna toL. virosa than toL. sativa/serriola. Such a discrepancy may be explained assuming that domestication ofL. sativa/serriola resulted in an increased selection pressure on unique DNA sequences as demonstrated by the RFLP data. Differential evolution of specific heterochromatin classes (and presumably of highly repetitive DNA classes), as revealed by chromosome banding techniques was not linked to domestication. Thus the disparity in conclusions about relationship (in terms of genetic similarity) as based on the different experimental approaches reflects a non-parallel evolution of highly repetitive vs. unique DNA classes.     

Intergeneric relationships ofLolium,Festuca, andVulpia (Poaceae) and their phylogeny

Morphological and seed protein analyses of 26 species of the generaLolium, Festuca andVulpia confirmed their close systematic affinities. Six inflorescence characters readily differentiatedFestuca fromLolium. Protein similarities betweenFestuca of sect.Bovinae and cross-pollinated species ofLolium, coupled with cytogenetic and crossability data, substantiate that they should be united into one genus.Vulpia had phenetic similarities with sect.Scariosae, Montanae andOvinae ofFestuca. Lolium, Festuca, andVulpia are most likely derived from a common ancestral form which was close toFestuca pratensis andLolium perenne.     

Vergleichende cytotaxonomische Untersuchungen anRanunculus seguieri und der Artengruppe desR. alpestris (Ranunculaceae)

Morphological and cytological investigations as well as crossing experiments were carried out withRanunculus seguieri Vill. and 4 species of theRanunculus alpestris L. group (R. alpestris L.,R. traunfellneri Hoppe,R. bilobus Bertol.,R. crenatus Waldst. & Kit.). ForR. seguieri andR. alpestris, localities and distribution are given in addition to extensive diagnoses. A key to the species includes morphological characteristics and distribution data forR. traunfellneri, R. bilobus, andR. crenatus. New diagnostic characters are described. Crossing experiments betweenR. seguieri and the species of theR. alpestris group were unsuccessful. All 5 species have a chromosome number of 2n = 16, the record forR. bilobus is new. There is no statistically significant difference between the karyotypes ofR. seguieri andR. alpestris s. str. Nevertheless, according to morphological evidence and crossing experiments,R. seguieri is not closely related to theR. alpestris group.

     

Molecular phylogeny and phylogeography ofLupinus (Leguminosae) inferred from nucleotide sequences of therbcL gene and ITS 1 + 2 regions of rDNA

Total DNA was extracted from 55 species of theLeguminosae (including 29 species ofLupinus). The chloroplast generbcL and the ITS 1 + 2 regions of nuclear RNA genes were amplified by polymerase chain reaction (PCR) and sequenced directly. The sequences obtained were evaluated with character state (Maximum Parsimony) and distance methods (Neighbour Joining). Phylogenetic trees obtained with both data sets and methods are mostly congruent.Genisteae andCrotalarieae are sister groups and share ancestry with theThermopsideae/Podalyrieae. The genusLupinus, which forms a monophyletic clade within theGenisteae, shows a distinct Old-New World disjunction and appears to be divided into several more or less distinct groups: (1) The species from the eastern part of South America. (2) The homogeneous rough-seeded group (Scabrispermae) of the Old World species which is well distinguished from the smooth-seeded group (Malacospermae). (3) Within the rather heterogeneous smooth-seeded lupins a smaller subgroup withL. angustifolius, L. hispanicus andL. luteus is recognized. (4) Also separated are North American lupins and South American species with a western distribution. Genetic distances imply that the genusLupinus evolved during the last 12–14 million years, ruling out the hypothesis that the present Old-New World disjunction can be interpreted as a result of the continental drift. The genetic data suggest an origin in the Old World and an independant colonisation of the Eastern parts of South America as opposed to North America and the Western parts of South America.     

Comparative spikelet morphology ofOryza schlechteri Pilger and related species ofLeersia andOryza (Poaceae)

Oryza schlechteri has only recently been rediscovered in Papua New Guinea. This species is quite distinct from otherOryza species but resembles closely related genusLeersia in its small panicle and profusely spreading habit. On the basis of spikelet morphology that has previously been used to distinguishLeersia andOryza species,O. schlechteri was compared withO. brachyantha. O. longiglumis, O. sativa, L. perrieri, L. tisseranti, andL. hexandra to clarify its generic position. Results corroborate that the presence of a sterile lemma and a striated spikelet epidermal (abaxial) surface lacking siliceous triads inO. schlechteri allies this species with otherOryza species rather than withLeersia.     

A second spontaneous hybrid in the genusLeucaena (Leguminosae, Mimosoideae)

The detection of hybridity inLeucaena is discussed in relation to: (i) traditional criteria, (ii) molecular criteria and (iii) models to predict hybrid leaf morphology. Morphological, geographical and molecular evidence for the occurrence of interspecific hybrids betweenL. leucocephala andL. diversifolia in south-central Mexico, northern Guatemala, Jamaica, Dominican Republic, the Philippines and Papua New Guinea is presented. Predicted mean hybrid leaf trait values calculated from parent material are compared with data from putative hybrids and shown to be similar. The origin of these hybrids is discussed and shown to be the result of artificial sympatry resulting from indigenous, and recent exotic, domestication of the parent species. The hybrid is described asL. ×spontanea.     

Responses of four arid zone grass species from varying habitats to drought stress

The effects of 4 or 8 drought cycles on four grass species,Cenchrus pennisetiformis, Leptochloa fusca, Panicum turgidum, andPennisetum divisum were assessed in a pot experiment. There were significant differences between the species in biomass production under water stress.C. pennisetiformis andP. turgidum produced significantly greater fresh and dry matter thanP. divisum and especially thanL. fusca. L. fusca had the lowest andP. divisum highest osmotic potentials compared with the other species after the completion of 4 or 8 drought cycles. Osmotic adjustment (difference between osmotic potential of droughted/rehydrated plants and control plants) was highest inL. fusca. The stomatal conductance was significantly decreased with increased drought stress inC. pennisetiformis. The elasticity ofC. pennisetiformis, P. turgidum andP. divisum increased with increase in number of drought cycles, whereas that ofL. fusca remained unchanged.L. fusca andP. turgidum had the lowest leaf hydration of all species after 8 drought cycles. The chlorophyllsa andb in all species remained unaffected by drought treatments. The proline content ofC. pennisetiformis andL. fusca increased significantly with increased drought stress, whereas that ofP. turgidum remained unaffected after 4 or 8 drought cycles.L. fusca synthesized great amount of leaf soluble proteins during 8 drought cycles, whereasP. divisum had low protein content after 4 drought cycles. The protein contents ofC. pennisetiformis andP. turgidum remained unaffected after 8 drought cycles. The leaf epicuticular wax ofL. fusca increased consistently with increased drought stress, but leaf wax ofP. divisum increased only at the highest drought stress and that ofC. pennisetiformis andP. turgidum increased after 4 drought cycles. On the basis of these results it was established thatC. pennisetiformis andP. turgidum were the most tolerant,P. divisum intermediate, andL. fusca the most sensitive to drought stress. The osmotic adjustment did not positively correlate with the degree of drought resistance.     

Phenetic investigation of non-nodulating African species ofAcacia (Leguminosae) using morphological and molecular markers

Morphological and RAPD markers were used to assess the relationships among nodulating and non-nodulating species of AfricanAcacia. Non-nodulating species of AfricanAcacia are only found within subg.Aculeiferum sect.Monacanthea. African species of sect.Monacanthea examined were found to form a group distinct from the other African species examined on a morphological and molecular basis. All lack the ability to nodulate, suggesting that non-nodulation may be used as a taxonomic tool. The species of sect.Aculeiferum were separated by RAPD and morphological analysis into two groups depending on whether they were armed with prickles in pairs and/or prickles in threes, or solitary. A third group of species was identified within sect.Acacia: further subdivision of this group was achieved into subsectt.Pluriseriae andUniseriae. The position ofA. albida relative to other AfricanAcacia species was found to be distinct but not totally independent of the genus. The partitioning and distribution of the genetic variability within the genus is further elucidated by the RAPD analysis of populations ofAcacia species. A population analysis ofA. polyacantha demonstrated geographical and site-specific variation.     

Morphological and molecular comparisons ofCampsis grandiflora andC. radicans (Bignoniaceae), an eastern Asian and eastern North American vicariad species pair

Morphological and molecular comparisons were made forCampsis grandiflora (Thunb.)K. Schumann (Bignoniaceae) from eastern Asia andC. radicans (L.)Seemann from eastern North America. Chloroplast DNA (cpDNA) variation was surveyed with 20 restriction endonucleases. The cpDNA divergence between the two vicariad species was 2.44%, which is the highest reported among North Temperate disjunct taxa and one of the highest reported for infrageneric taxa. Detailed morphological comparisons also suggest a high level of divergence. Cluster analyses based on 22 morphological characters and 39 OTUs revealed two distinct groups corresponding with the two species. The average taxonomic distance between the two species was 1.806. Analyses of variance (ANOVA) revealed that 12 of the 18 quantitative characters differed significantly ( 0.01) betweenC. grandiflora andC. radicans. Divergence time based on cpDNA data was estimated as 24.4 million years. The Bering land bridge hypothesis was favored over the North Atlantic land bridge hypothesis based on the estimated divergence time and the geological history of the North Temperate region. The high levels of morphological and cpDNA divergence are not consistent with morphological stasis, which has been proposed as a common mode of evolution for North Temperate disjunct taxa.     

Allozyme divergence and evolution in the genusLens

The genusLens includes 5 taxonomic species:L. culinaris is cultivated andL. orientalis, L. odemensis, L. ervoides, andL. nigricans are wild. All the species are annual and almost exlusively selfers. The wild lentils are distributed over a large geographical area and form small disjunct populations which are composed of a small number of plants. 67Lens populations were assayed electrophoretically for 9 enzyme systems; 15 enzymic genes with 37 alleles were identified. The genetic distances (D) measured between the pairs of populations indicated a significantly greater similarity between populations belonging to the same taxonomic species. Assuming the populations represent a random sample of the variability in each of the species the genetic distances (D) between the 5 taxa were calculated. The shortest genetic distance was found betweenL. orientalis andL. culinaris. Another significant feature of the data is the apparent isolation ofL. nigricans from the other 4 species. The genetic distances between theLens species are compared to the patterns of crossability barriers between them.     

Pollen morphology inParis and its related genera

Pollen grains ofParis, Medeola, andScoliopus have been examined with light, scanning electron, and transmission electron microscopies.P. quadrifolia L.,P. verticillata Bieb.,P. delavayi Franch. andP. polyphylla Smith have monosulcate pollen characterized by psilate, foveolate or reticulate exine sculpture. In contrast to the former species,P. japonica (Fr. et Sav.) Franch. andP. tetraphylla A. Gray have monosulcate pollen with gemmate (rarely rugulate) exine.Medeola has monosulcate pollen with reticulate exine that is distinct from that ofParis. Scoliopus has monosulcate pollen characterized by a peculiar reticulate exine pattern. The palynological evidence suggests thatParis andTrillium are closely related to each other, andMedeola andScoliopus should be separated fromParis andTrillium.     

Coexistence of predatoryMuscina stabulans andophyra aenescens [Dipt.: Muscidae] with dipterous prey in poultry manure

Significant relationships between the emergence of adult predatory species,Ophyra aenescens (Wiedemann) andMuscina stabulans (Fallen) and prey,Musca domestica L.,Fannia femoralis (Stein) andF. canicularis (L.) existed on 10 poultry ranches in southern California. Predatory species may function in natural control at intervals throughout the year, especially on predominant prey,F. femoralis andF. canicularis. Dipterous species, especiallyO. aenescens andF. femoralis, were only moderately influenced by the surface area of breeding habitat available. Accuracy in the measurement of population trends in Diptere may be enhanced by increasing the number of samples at any given ranch rather than the addition of study ranches. Parasitism, ranging from 22.6–37.8% of collected puparia, involved principally 3 species,Muscidifurax zaraptor Kogan & Legner,M. raptor Girault & Sanders andSpalangia cameroni Perkins.      

Distribution of parental DNA markers inEncelia virginensis (Asteraceae: Heliantheae), a diploid species of putative hybrid origin

Morphological, geographical and ecological evidence suggests thatEncelia virginensis is a true-breeding diploid species derived from hybrids ofE. actoni andE. frutescens. To test this hypothesis, we examined the chloroplast and nuclear DNA of severalEncelia species. PCR amplification targeted three separate regions of chloroplast DNA:trnK-2621/trnK-11,rbcL/ORF106, andpsbA3/TrnI-51, which amplify 2600bp, 3300bp and 3200bp fragments respectively. Restriction fragment analysis of chloroplast DNA revealed no variation that could be used to discriminate between the parent species. A RAPD analysis using 109 dekamer primers was used to analyze the nuclear genome.Encelia actoni andE. frutescens were distinguished by several high-frequency RAPD markers. In populations ofE. virginensis, these markers were detected in varying proportions, and no unique markers were found. Evidence from the nuclear genome supports the hypothesis thatE. virginensis is of hybrid origin. ThatE. virginensis may have arisen by normal divergent speciation followed by later introgression remains a possibility, however, and is not formally ruled out here. Diploid hybrid speciation inEncelia differs from other documented cases in that there are no discernible chromosome differences between the species, and all interspecific hybrids are fully fertile. In addition, apparent ecological selection against backcross progeny provides an external barrier to reproduction between F₁ progeny and the parental species. These characteristics suggest that hybrid speciation inEncelia may represent an alternative model for homoploid hybrid speciation involving external reproductive barriers. In particular, this may be the case for other proposed diploid hybrid taxa that also exhibit little chromosomal differentiation and have fertile F₁s.     

Presence of different arbuscular mycorrhizal infection patterns in roots of<Emphasis Type="Italic"> Lotus glaber</Emphasis> plants growing in the Salado River basin

Morphological types of arbuscular mycorrhizal (AM) fungi associated with Lotus glaber in sodic soils of the Salado River basin were studied. At least eight colonization patterns (IP) of AM fungi in roots of L. glaber were observed after 30 plants were analyzed. Arum- and Paris-type infection were found in the same plant species. This result supports the idea that AM morphology is not solely under plant control, but is also influenced by fungal identity. One infection pattern, presumably corresponding to Glomus intraradices, and a second, possibly assignable to Glomus tenue, were the most commonly found. Our results reinforce previous suggestions that G. intraradices is well adapted to sodic-saline conditions and may play a role in the resistance of L. glaber to these soils.     

Labiatae Novae Iranicae

Four new species are described:Lavandula sublepidota, from S. Iran (related toL. coronopifolia),Phlomis pachyphylla from S. Iran, Prov. Fars (nearPh. polioxantha),Satureia isophylla from N. Iran, Prov. Mazanderan (close toS. mutica andS. intermedia), andThymbra neurophylla from Iraqi Kurdistan (distinct forth species of the genus).

Florae Iranicae praecursores 1–4.     

A survey of C-band patterns in chromosomes ofLilium (Liliaceae)

C-band patterns are described for 20Lilium spp. distributed across six sections. All species have a similar basic karyotype (n = 12) but C-bands differ markedly between them. The patterns are characterized by a dispersed scattering of thin intercalary bands as well as centric and NOR bands. Only one species,L. canadense, shows a clear equilocal pattern with intercalary C-bands occurring proximally in all of the longer chromosome arms. Comparing species, similar patterns are revealed forL. regale andL. sulphureum, forL. formosanum andL. longiflorum (all in sect.Leucolirion) and to a lesser extent forL. hansonii, L. martagon, andL. tsingtauense (sect.Martagon). The pattern forL. henryi (previously classed in sect.Sinomartagon) matches those ofL. regale andL. sulphureum quite well and its transfer to sect.Leucolirion is proposed. This is consistent with results from interspecies hybrids betweenL. henryi andL. regale (and related species) which are reportedly fertile. No other clear similarities in C-band patterns were seen across species. It seems that C-band patterns change rapidly inLilium and hence their usefulness in classification will be restricted to identifying closely related species.Dedicated to Prof.D. G. Catcheside on the 80th anniversary of his birth.     

Cytotaxonomy of the genusPolystichum in the Western Himalayas

Cytotaxonomical investigations on 11 species from the Western Himalayas show that 8 are diploid and the remaining tetraploid. The numbers for three species are new.P. prescottianum (2x) andP. thomsoni (4x), andP. stimulans (2x) andP. acanthophyllum (4x) are closely related but distinct species pairs.     

Patterns of intraspecific variation inAntennaria alborosea,A. corymbosa,A. marginata,A. microphylla,A. parvifolia,andA. umbrinella (Asteraceae:Inuleae)

Patterns of intraspecific variation were examined inAntennaria alborosea A. E. Porsild,A. corymbosa E. Nels,A. marginata Greene,A. microphylla Rydb.,A. parvifolia Nutt., andA. umbrinella Rydb. AlthoughA. alborosea was initially considered arctic in distribution, it became apparent that a southern montane element also exists. Our results suggest that morphological differences between arctic and southern montane specimens represent clinal variation. The additional morphological data for specimens that occur more than 1,500 km south of the species' range as it was initially described result in a better understanding of this once presumed arctic taxon. Morphological variation in the dioecious speciesA. corymbosa, A. marginata, A. microphylla, A. parvifolia, andA. umbrinella was greater between the genders than was geographic variation within each gender. These results demonstrate that both pistillate and staminate specimens must be examined in dioecious species ofAntennaria if morphological variation in the respective species is to be fully understood. Character size or number of broadly distributed species (A. microphylla andA. parvifolia) generally decreased with increasing longitude, whereas characters of species with more restricted distributions (A. alborosea, A. corymbosa, andA. marginata) generally increased in size or number with increasing latitude or longitude.Antennaria umbrinella was an exception in this respect.