The so‐called primitive genera of Genisteae (Fabaceae): systematic and phyletic considerations based on karyological data

A karyological analysis of the so‐called primitive genera of Genisteae has shown that they have a relatively homogeneous chromosome complement: all species tend to have a somatic chromosome number 2n = 48, which can increase to 2n = 52, presumably as a result of hyperaneuploidy. Karyological data suggest that Argyrocytisus, Cytisophyllum and Petteria may be considered as distinct genera rather than being assigned to Cytisus, with 2n = 52 for the first of these and 2n = 50 for the other two genera. They may be interpreted as relict monotypic genera as a result of the presence of a stabilized aneuploidy. Karyological characters exclude a recent origin of Genisteae from Thermopsideae. On the contrary, they are consistent with the hypothesis that Genisteae and Thermopsideae are independently derived from a basic papilionoid stock, of which present day Sophoreae are the remainder. At least two lines would lead from Sophoreae to the taxa of the ‘genistoid alliance’, one to Thermopsideae and the other ‘podalyrioid alliances’ (Podalyrieae and Mirbelieae), with the prevailing basic number of x = 9, and the other to Genisteae, with a basic number of x = 12 persisting in some present day genera, including Cytisus s.l. From this lineage, a wide range of secondary basic numbers has been formed, mostly by descending aneuploidy. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 232–248.     

A phylogeny of the chloroplast gene rbcL in the Leguminosae: taxonomic correlations and insights into the evolution of nodulation

Phylogenetic analysis of the chloroplast-encoded rbcL gene in Leguminosae are consistent with previous hypotheses in suggesting that the family as a whole is monophyletic, but that only two of its three subfamilies are natural. The earliest dichotomies in the family appear to have involved tribes Cercideae or Cassieae (subtribe Dialiinae), followed by Detarieae/ Macrolobieae, all of which are members of subfamily Caesalpinioideae. The remainder of the family is divided into two clades: (1) Mimosoideae and the caesalpinioid tribes Caeasalpinieae and Cassieae (subtribes Ceratoniinae and Cassiinae); (2) Papilionoideae. Basal groups within Papilionoideae are, as expected, elements of the grade tribes Sophoreae and Swartzieae. Major clades within Papilionoideae include: (1) a Genistoid Alliance comprising Genisteae, Crotalarieae, Podalyrieae, Thermopsideae, Euchresteae, and also some Sophoreae; (2) a clade marked by the absence of one copy of the chloroplast inverted repeat, with which are associated Robinieae. Loteae, and some Sophoreae; (3) Phaseoleae, Desmodieae. Psoraleeae, and most Millettieae, a group also marked by presence of pseudoracemose inflorescences; and (4) a well-supported clade comprising Aeschynomeneae, Adesmieae, and some Dalbergieae. Nodulation is most parsimoniously optimized on the rbcL strict consensus tree as three parallel gains, occurring in Papilionoideae, the caesalpioioid ancestors of Mimosoideae, and in the genus Chamaecrista (Caesalpinieae: Cassieae).     

Quinolizidine alkaloids as systematic markers of the papilionoideae

The structural characteristics of quinolizidine alkaloids have been quantified on the basis of their biogenetic origins. Such data have been used to determine evolutionary advancement parameters for genera of the Papilionoideae. These analyses indicate a correlation of the structural characters of alkaloids with ecogeography and morphology of the genera, suggesting an adaptive radiation of the taxa from tropical Africa in three major directions along a southern temperate “pre-cytisine” route, a northern temperatre “pre-cytisine-cytisine” route and a tropical “ormosanine” route with a temperate cytisine branch. The data also suggest the division of the Sophoreae into three chemical groups giving rise to Liparieae, Genisteae, Thermopsideae and Bossiaeeae. These concepts are embodied in a tentative phyletic classification of all genera of the Papilionoideae which contain quinolizidine alkaloids.     

Alkaloid distribution in some species of the Papilionaceous tribes Thermopsideae and Genisteae

Alkaloid profiles of extracts of fourteen papilionaceous species in the tribes Thermopsideae and Genisteae (sensu Polhill) have been determined. Altogether, eighteen quinolizidine alkaloids, representative of four structural groups, and one dipiperidine alkaloid were identified among the legumes studied. The chemotaxonomic implications of these data are discussed.     

Serotaxonomy and systematic relationships of the genus Adenocarpus (Genisteae – Fabaceae)

The storage proteins extracted from the seeds of five species of the genus Adenocarpus and of four subspecies or varieties of the type species A. complicates were studied serologically and compared with the homologous proteins extracted from 20 other species of Genisteae and allied tribes. All taxa of Adenocarpus reacted strongly with an antiserum to A. complicates ssp. complicates; only slight differences were observed between subspecies and between species. Adenocarpus was shown to be isolated from all other Genisteae. However, it was more similar to the Genisteae than to species of other tribes. The highest similarity toward Adenocarpus was presented by Argyrolobium , by some primitive unspecialized genera (Laburnum, Cytisophyllum) , and by such isolated genera as Cytisanthus, Spartium, Echinospartum. Retama and Lupinus were most dissimilar. It is concluded that Adenocarpus , a very uniform and isolated genus, belongs to the Genisteae; it shares many characters with the bulk of primitive genera of the Genisteae; its evolutionary line is independent from all others, but is not very distant from that of Argyrolobium.     

A phylogeny of Thermopsideae (Leguminosae: Papilionoideae) inferred from nuclear ribosomal internal transcribed spacer (ITS) sequences

Based on nuclear ribosomal DNA internal transcribed spacer (ITS) sequences, Thermopsideae is phylogenetically studied within a genistoid background. Analysis reveals that the tribe is not supported as a monophyletic group. Some species of Sophora s.s are nested within it. The central Asian desert Ammopiptanthus forms an isolated clade but is relatively remote to other Thermopsideae members. Piptanthus , Anagyris , Baptisia , and Thermopsis are clustered together into a robust clade. We hence propose that the tribe could either be reduced to just the four 'core genera' with Ammopiptanthus excluded, or, as an alternative, that Thermopsideae could become part of a new Sophoreae s.s. if it is re-circumscribed in the future. Both Piptanthus and Baptisia appear as monophyletic. The genus Anagyris is closer to some east Asian Thermopsis species than to Piptanthus . The east Asian and North American disjunct Thermopsis is not monophyletic. The ITS results suggest a geographical division between the Old World and New World Thermopsis . The east Asian species are clustered with Piptanthus and Anagyris , whereas the North American species are allied to Baptisia . Nonetheless, the only two north-eastern east Asian native Thermopsis species appear to be more related to the North American group than to the east Asian one. The related biogeographical significance has therefore been additionally discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 365–373.     

Sero-systematic and taximetric studies on the Phaseoleae (Fabaceae) and related tribes

VIVIANI, T., CONTE, L., CRISTOFOLINI, G. & SPERANZA, M., 1991. Sero-systematic and taximetric studies on the Phaseoleae (Fabaceae) and related tribes. One hundred and fifty-two species of Phaseoleae, Desmodieae, Indigofereae, Millettieae, Sophoreae and Swartzieae were subjected to a taximetric study. The serological relationships were analysed using 37 antigenic systems and 12 antisera upon a representative sample from the six tribes. The comparative use of taximetric and serological data allowed the following conclusions to be drawn: (1) the tribe Phaseoleae is not a natural taxon; the primitive complex Diocleinae-Glycininae, the more derived Phaseolinae, and the Cajaninae deserve tribal rank; the position of the remaining subtribes requires further examination. (2) The Desmodieae are related to the Phaseoleae, whereas the Indigofereae have little relation to them. (3) The Baphia -group in the Sophoreae is possibly related to the Phaseoleae ancestors, while the other Sophoreae belong to different phyletic lines. (4) Swartzia is morphologically related to the Sophoreae, but does not belong to the same natural group. (5) Sophoreae, Millettieae and Phaseoleae are three heterogeneous aggregates representing stages of increasing specialization, rather than elements of a single phyletic line.     

Pollen morphology in relation to pollinators in Papilionoideae (Leguminosae)

The surface sculpturing of the pollen of some species of the taxonomically widely separated genera Harpalyce (tribe Brongniartieae), Camoensia (tribe Sophoreae), Millettia (tribe Tephrosieae), and of the monotypic Dahlstedtia (tribe Tephrosieae) which have large red or white flowers adapted for pollination by birds or bats, is coarsely rugulate or verrucate. Related taxa with small insect pollinated flowers have pollen with simple reticulate or perforate surface sculpturing. The exine stratification of Alexa and Castanospermum (tribe Sophoreae), genera with large red bird-flowers, is complex with a layer of tectal columellae and differs from that of other genera in the tribe Sophoreae which have a normal pollen wall structure. These modifications of pollen structure and sculpture appear to be the result of convergent evolution and a secondary adaptation to pollination. The taxonomic and functional significance of the observations are briefly discussed.     

Serological contributions to the systematics of Ulex (Genisteae – Fabaceae) and allied genera

Seven Ulex species were compared serologically with 21 species of related genera. Seed proteins were employed to immunize rabbits; immune sera to 17 species were produced. The presence or absence of differential characters (spurs) was detected in immunodiffusion experiments and recorded in a square matrix for each antiserum. The interspecific correlation was then computed, and an ordination on principal components was obtained.
The results suggested that: (1) Ulex L. should include sect. Pseudogenista (= genus Nepa Webb) and sect. Slauracanthus (= genus Stauracanthus Link); (2) sect. Stauracanihus is closer to sect. Ulex than to sect. Pseudogenista; (3) Ulex is markedly dissimilar from any other genus, its highest similarity being to Genista sect. Phyllobotrys and to Chamaespartium; (4) Ulex behaves like other derived genera and can be regarded as an extremely derived form within the Genista evolutionary line; (5) Echinospartum seems a primitive genus, it is not closely related to Ulex but rather to Cytisanthus and allied genera. There is a good agreement between these results and morphological, phytogeographical and caryological evidence. A general outline of evolutionary relationships within the Genisteae is presented.     

Diversity of rhizobia nodulating wild shrubs of Sicily and some neighbouring islands

Legume shrubs have great potential for rehabilitation of semi-arid degraded soils in Mediterranean ecosystems as they establish mutualistic symbiosis with N-fixing rhizobia. Eighty-eight symbiotic rhizobia were isolated from seven wild legume shrubs native of Sicily (Southern Italy) and grouped in operational taxonomic units (OTU) by analysis of the ribosomal internal transcribed spacer (ITS) polymorphism. Partial sequencing of 16S rRNA gene of representative isolates of each OTU revealed that most Genisteae symbionts are related to Bradyrhizobium canariense, B. japonicum and B. elkanii. Teline monspessulana was the only Genistea nodulated by Mesorhizobium strains, and Anagyris foetida (Thermopsideae) was promiscuosly nodulated by Rhizobium, Mesorhizobium, Agrobacterium and Bradyrhizobium strains. Analysis of the nodulation gene nodA assigned most Mediterranean Genisteae bradyrhizobia to clade II but also to clades IV, I and III, which included, so far, sequences of (sub)tropical and Australian isolates. The high diversity and low host specificity observed in most wild legumes isolates suggest that preferential associations may establish in the field depending on differences in the benefits conferred to the host and on competition ability. Once identified, these beneficial symbiosis can be exploited for rehabilitation of arid, low productive and human-impacted soils of the Mediterranean countries.     

A new genus segregated from Thermopsis (Fabaceae: Papilionoideae): Vuralia

A relict endemic species originating from Turkey, Thermopsis turcica, was excluded from Thermopsis in consideration of morphologic studies based on its flowers and fruits as well as micromorphological features that were obtained by scanning electron microscopy (SEM) and data from supporting molecular analyses internal transcribed spacer (ITS). Additionally, phylogenetic analyses (maximum parsimony and neighbour-joining) were conducted using PAUP 4.0 software. These analyses supported a new taxonomical position for the Turkish species. SEM and morphological studies indicated that the species has unique features that have not been observed in other Thermopsideae species. The unique features and characteristics are comprehensively illustrated here. For confirmation of our purpose, the chromosome number and karyotype of V. turcica were determined via the KAMERAM software program. As a result, all detailed examinations and comparisons of Vuralia and related genera suggest that Vuralia is a new monotypic genus within Thermopsideae, and a new genus and combination nova were established for this Turkish species.     

Genetic characterization of wild leguminous nodular bacteria living in the South Urals

Genetic diversity and phylogeny of rhizobia that nodulate 18 species of wild-growing bean plants of South Urals from 8 genera belonging to 4 tribes (Loteae, Genisteae, Galegeaev and Hedysareae) was studied. It was demonstrated that for the wild-growing plants of Galegeae and Hedysareae tribes symbiotic interaction with various strains of nodule bacteria that closely related to bacteria of Mesorhizobium sp. was typical of the plants of Genisteae tribe--to bacteria of Bradyrhizobium sp. In the nodules of Lortus ucrainicus from Loteae tribe we have found a rhizobium that is closely related to the bacteria of Mesorhizobium sp., and at Coronilla varia rhizobia strains obtained by us were close by sequence of a 16S pRNA gene to Rhizobium sp. In the nodules of some kinds of the investigated plants we found also minor species of a rhizobia, which structure is under the great influence of conditions of the host plant growth.     

A serological contribution to the systematics of the genusLupinus (Fabaceae)

The immunological comparison of the seed reserve proteins suggests thatLupinus is a natural genus, the American and the Euro-African species belonging to the same stock. Among the Lupines of the Old World, the smooth-seeded and the rough-seeded species from two natural segregates. The genusLupinus is serologically related to the rest of theGenisteae and to the AsiaticSophoreae rather than to AmericanSophoreae andThermopsideae. The data suggest thatLupinus may have originated with the remainder of theGenisteae from primitiveSophoreae of temperate-subtropical Asia. America and the Mediterranean-African region are regarded as secondary centres of speciation.     

The Systematic and Ecological Significance of Anatomical Characters of Leaves in the Tribe Thermopsideae (Fabaceae)

Anatomical characters of leaves of Chinese species of the tribe Thermopsideae, including 3 genera and 13 species, were compared. The results show that the differentiation of mesophyll cells, stomatal types, stomatal distribution and density, shape of epidermal cell, trichome shape and density, cuticular membrane and waxy ornamentation have important systematic and ecological significance among the genera of the tribe. With regard to leaf architecture, pollen morphology and external morphology studies, we suggest that Ammopiptanthus represents an isolated and primitive genus in the tribe and Piptanthus is closely related to Thermopsis. Although different vein-endings, stomatal types and trichomes indicate systematic differences, other characters, such as mesophyll type, stomatal distribution, trichome density etc. are the result of ecological adaptation among the genera. Results of the present study confirm our previous conclusion that two evolutionary trends exist in the tribe, the first towards xerophilization and the second towards cold-adaptation They are caused by the desertification of the central Asian area and the heave of intermountainous plateau. Ecological isolation may be a main evolutary motivation among the taxa in the tribe. We have obs'erved that stomatal density and size in the xeromorphic genus, Ammopipianthus, are not simply a result of the general trend in xerophytes to have high stomatal density correlating with the relatively small size. On the contrary, stomata in this genus are larger than those in the mesophytic genus, Piptanthus, and the density on the lower surface is much less than in the latter. However, stomatal density on the upper surface follows the trend in concordance with increasing aridity and altitute.     

Genetic variation and phylogeography of Stauracanthus (Fabaceae, Genisteae) from the Iberian Peninsula and northern Morocco assessed by chloroplast microsatellite (cpSSR) markers

The tribe Genisteae includes genera of great ecological importance in Mediterranean countries because they are dominant elements of many plant communities. Genetic variation and diversification patterns in Stauracanthus (Genisteae) provide information relevant for the study of the processes of diversification in relation to the environmental history of the western Mediterranean. Nineteen populations of S. boivinii and S. genistoides were assessed by 11 chloroplast microsatellite markers, revealing 44 haplotypes. Both species had different haplotypes and contrasting patterns of karyological, morphological, and genetic variation. In the minimum spanning tree of the haplotypes, AMOVA analysis, and nested clade analysis, S. boivinii had high levels of differentiation and restricted gene flow among populations. Allopatric differentiation occurred between the Moroccan and Iberian populations of S. genistoides, although S. genistoides subsp. spectabilis and subsp. vicentinus had high levels of differentiation among populations (F(ST)), whereas S. genistoides subsp. genistoides had a low F(ST). Genetic patterns are discussed in relation to the Messinian salinity crisis (MSC): hard conditions drove plants to refuge habitats along the Atlantic coast and higher altitude areas in the Moroccan mountains (S. genistoides subsp. spectabilis and S. boivinii). After the MSC, S. boivinii underwent polyploidization and expansion, whereas S. genistoides expanded and continued diversifying into S. genistoides subspp. genistoides and vicentinus.     

CHROMOSOME NUMBERS IN THE LEGUMINOSAE II: AFRICAN SPECIES,INCLUDING PHYLETIC INTERPRETATIONS

Turner , B. L., and O. S. Fearing . (U. Texas, Austin.) Chromosome numbers in the Leguminosae. II. African species, including phyletic interpretations. Amer. Jour. Bot. 46(1) : 49-57. Illus. 1959.—Chromosome numbers for 30 African legume species have been reported. These include first reports for 28 taxa, including 12 genera (Bolusanthus, Calpurnia, Melolobium, Lessertia, Sulherlandia, Colophospermum, Guibourtia, Burkea, Julbernardia, Schotia, Piliostigma and Swartzia). The counts are discussed with respect to those previously reported for related groups, and this chromosomal information was used to construct hypothetical phyletic lines at the tribal level within the subfamilies Papilionoideae and Caesalpinioideae. A phyletic scheme for the Leguminosae (excluding the Mimosoideae) based on this evidence from chromosome studies is presented. Notable departures from previously suggested phyletic treatments include: (1) Suggestion for inclusion of genera of the Galegeae and Hedysareae with base numbers of x = 10 and 11 with the Phaseoleae and Dalbergieae. (2) Derivation of the Papilionoideae through caesalpinoid prototypes, possibly from Swartzia-like ancestors. (3) Recognition of several very old chromosomal lines stemming from the subfamily Caesalpinioideae, and the suggestion that parts of the tribes Sclerolobieae, Cynometreae, Swartzieae and Sophoreae are, perhaps, more closely related to each other and to the Papilionoideae than they are to the remaining caesalpinoid tribal lines.     

中国食线虫真菌的种类

对我国24个省市,涉及40余种植物的300多个土样的调查,分离出食线虫真菌近千个分离物,共鉴定出食线虫真菌29属57种,包括捕食线虫真菌4属25种,其中Arthrobotrys arthrobotryoides, A. brochopaga, A. cladodes var. macroides, A. microspora,A. musiformis, A. straminicola, Monacrosporium ellipsosporinm, M. thaumasium, M. pravicollis,Stylopage grandis, Dactylella ramiformis, Monacrosporium qiuanum, Monacrosporium sinense等10种为国内新记录。内寄生真菌4属4个种、其中Catenaria angullulae, Harposporiumarcuatum, Nematoctonus leiosporus为国内新记录。定殖于大豆孢囊线虫真菌22属25种,其中Catenaria auxiliaris, Cylindrocarpon heteronema, C. willkommii, Fusariella bizzozeriana,Nematophthora sp., Phoma eupyrena, Phoma leveilei,Pithomyces sp,Trichocladium opacum.Verticillium chlamydosporium等10属种为国内新记录,其中Nematophthora, Fusariella,Trichocladium三属为国内新记录属,Rhizoctonia sp., Stachybotrys elegans, F.bizzozeriana,C.heteronema, C.willkommii等为首次发现定殖于大豆孢囊线虫孢囊上。对所鉴定的真菌进行了描述和讨论并制作了检索表。     

Cross-species amplification of soybean (Glycine max) simple sequence repeats (SSRs) within the genus and other legume genera: implications for the transferability of SSRs in plants

We investigated the transferability of 31 soybean (Glycine max) simple sequence repeat (SSR) loci to wild congeners and to other legume genera. Up to 65% of the soybean primer pairs amplified SSRs within Glycine, but frequently, the SSRs were short and interrupted compared with those of soybeans. Nevertheless, 85% of the loci were polymorphic within G. clandestina. Cross-species amplification outside of the genus was much lower (3%-13%), with polymorphism restricted to one primer pair, AG81. AG81 amplified loci in Glycine, Kennedia, and Vigna (Phaseoleae), Vicia (Vicieae), Trifolium (Trifolieae), and Lupinus (Genisteae) within the Papilionoideae, and in Albizia within the Mimosoideae. The primer conservation at AG81 may be explained by its apparent proximity to the seryl-tRNA synthetase gene. Interspecific differences in allele size at AG81 loci reflected repeat length variation within the SSR region and indels in the flanking region. Alleles of identical size with different underlying sequences (size homoplasy) were observed. Our findings and the emerging patterns in other plant studies suggest that in contrast to animals, successful cross-species amplification of SSRs in plants is largely restricted to congeners or closely related genera. Because mutations in both the SSR region and the flanking region contribute to variation in allele size among species, knowledge of DNA sequence is essential before SSR loci can be meaningfully used to address applied and evolutionary questions.      

The Variability of Organs Differentially Involved in Pollination, and Correlations of Traits in Genisteae (Leguminosae: Papilionoideae)

Size differences in organs involved to varying degrees in pollenexchange (standard and keel petals, calyx, pedicel and inflorescencestem diameter), together with correlations among these traits,were investigated at both the intraspecific and interspecificlevels in populations of 15 southern Spanish species of thepapilionoid legume tribe Genisteae. Characteristics less relatedto pollination (e.g. pedicel length and stem diameter) wereroughly twice as variable between individuals as petal or calyxdimensions. The coefficients of variation were in the range16–19%vs. 7–10%, respectively. Standard and keelpetal size varied in parallel, not only in conspecific individualsbut also across species and genera, suggesting a tight developmentalrelationship between these organs and a low potential for evolutionarymodification. The remaining characteristics covaried in differentways within and between species and thus appeared much lessmutually constrained (particularly so for calyx size). Multivariateanalyses of variance revealed significant population divergencein five out of six species. Overall, 28% of all variation inflower size occurred among populations and 72% within populations,but the proportion of variance accounted for by population membershipvaried extensively across species (from 1 to 64%) and was highestin two selfing annuals (Lupinus). Outcrossing perennials mayshow either high (Stauracanthus, Genista) or low (Cytisus) populationdifferentiation. In addition to contributing to the increasingamount of data on quantitative floral variation in plants, resultsof this study may help understand how pleiotropic effects limitthe evolutionary modification of some flower traits more thanothers in diverse plant groups. Copyright 2001 Annals of BotanyCompany Calyx, corolla, Genisteae, legumes, shrub, population divergence, Mediterranean, pollination, individual variation     

Systematic consideration of floral microcharacters of the South American genus Chrysolaena (Vernonieae,Asteraceae)

Floral microcharacters of the genus Chrysolaena H. Rob. (Vernonieae, Asteraceae) were analysed in detail for the first time in order to evaluate the taxonomic position of conflictive species in the group. The results were also compared with studies carried out in species of related genera. In addition to distinctive microcharacters previously studied in some species of the genus, other characters such as trichome types of the corolla, style, anthers and cypselae have been analysed for the first time. The presence of glandular apical appendage and cypselae are common characteristics among species Chrysolaena. In addition to these, this study shows that presence of glands on the style and corollas is another consistent characteristic in the genus. However, the absence of basal stylar node would not be a diagnostic character since this varies widely among species. The results indicate that most of the microcharacters of Chrysolaena analysed are quite consistent in the genus, but they are no more consistent than the pollen morphology (type 'C') and chromosome base number (x = 10). Until now, these last two features would be most useful for separating Chrysolaena from the related genera Lessingianthus and Lepidaploa. At species level, the results show that related species can be distinguished by the different combinations of floral microcharacters. The value of microcharacters could be increased if they are combined with other morphological, cytological, and palynological data.