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In most angiosperms, the endosperm develops before the embryo, but with harmony between the two structures until final seed formation. In an embryological study, we show that inbreeding depression causes disharmony in development of the two structures in two Leguminosae shrubs, Cytisus multiflorus and C. striatus. Our main objective was to test the causes of self- sterility in the two species by comparing the embryological development of the self seeds with that of cross seeds. In developing selfed seeds of C. multiflorus, the embryo reaches at most the globular stage and never forms mature seeds, while in C. striatus a few mature selfed seeds are formed. In both species, the main cause of abortion of developing selfed seeds is diminished endosperm development (low values of the ratio of endosperm to embryo), which triggers collapse of the endosperm and embryo. The results indicate that self-sterility in C. striatus is postzygotic because of strong, early inbreeding depression, while in C. multiflorus there exists a mixed pre- and postzygotic mechanism; the prezygotic mechanism causes rejection of some self-pollen tubes in the style/ovary, and the early inbreeding depression triggers abortion of fertilized ovules that escaped that action. Key words: abortion; Cytisus multiflorus; Cytisus striatus; embryogeny failures; inbreeding depression; late-acting self-incompatibility; Leguminosae; Mediterranean region; self-sterility. doi: 10.3732/ajb.0800332

This analysis goes beyond many phylogenies in exploring how phylogenetic structure imposed by morphology, ecology, and geography reveals useful evolutionary data. A comprehensive range of such diversity is evaluated within tribe Indigofereae and outgroups from sister tribes. A combined data set of 321 taxa (over one-third of the tribe) by 80 morphological characters, 833 aligned nuclear ribosomal ITS/5.8S sites, and an indel data set of 33 characters was subjected to parsimony analysis. Notable results include the Madagascan dry forest Disynstemon resolved as sister to tribe Indigofereae, and all species of the large genus Indigofera comprise just four main clades, each diagnosable by morphological synapomorphies and ecological and geographical predilections. These results suggest niche conservation (ecology) and dispersal limitation (geography) are important processes rendering signature shapes to the Indigofereae phylogeny in different biomes. Clades confined to temperate and succulent-rich biomes are more dispersal limited and have more geographical phylogenetic structure than those inhabiting tropical grass-rich vegetation. The African arid corridor, particularly the Namib center of endemism, harbors many of the oldest Indigofera lineages. A rates analysis of nucleotide substitutions confirms that the ages of the oldest crown clades are mostly younger than 16 Ma, implicating dispersal in explaining the worldwide distribution of the tribe. Key words: biogeography; character evolution: dispersal limitation; Fabaceae; Indigofera; Indigofereae; Leguminosae; molecular phylogeny; rates analysis.

The endangered Shivwits milkvetch, Astragalus ampullarioides, is a perennial, herbaceous plant. This Utah endemic was federally listed as endangered in 2001 because of its high habitat specificity and low numbers of individuals and populations. All habitat currently occupied by A. ampullarioides was designated as critical by the U.S. Fish and Wildlife Service in 2006 as a result of conservation litigation. We used AFLP markers to assess genetic differentiation among the seven extant populations and quantified genetic diversity in each. Six different AFLP markers resulted in 217 unambiguous polymorphic loci. We used multiple methods to examine any changes in population genetic structure in this species over time. Results indicate that A. ampullarioides had much higher gene flow among populations in the past, but has since fragmented into regional genetic units. These regions further fragmented genetically, and extant populations have differentiated through genetic drift. Populations had low levels of gene flow, even between geographically close populations. Rapid urban development reduces gene flow among regions and encroaches on populations of A. ampullarioides and remaining patches of unoccupied habitat. The genetic makeup of each of the extant populations should be carefully considered in management decisions such as population establishment or augmentation. Key words: AFLP; Astragalus ampullarioides; endemic; Fabaceae; population genetics; soil specific; Washington County, Utah; Utah; Zion National Park.

Prosopis represents a valuable forest resource in arid and semiarid regions. Management of promising species requires information about genetic parameters, mainly the heritability ([h.sup.2]) of quantitative profitable traits. This parameter is traditionally estimated from progeny tests or half-sib analysis conducted in experimental stands. Such an approach estimates [h.sup.2] from the ratio of between-family/total phenotypic variance. These analyses are difficult to apply to natural populations of species with a long life cycle, overlapping generations, and a mixed mating system, without genealogical information. A promising alternative is the use of molecular marker information to infer relatedness between individuals and to estimate [h.sup.2] from the regression of phenotypic similarity on inferred relatedness. In the current study we compared [h.sup.2] of 13 quantitative traits estimated by these two methods in an experimental stand of P. alba, where genealogical information was available. We inferred pairwise relatedness by Ritland's method using six microsatellite loci. Relatedness and heritability estimates from molecular information were highly correlated to the values obtained from genealogical data. Although Ritland's method yields lower [h.sup.2] estimates and tends to overestimate genetic correlations between traits, this approach is useful to predict the expected relative gain of different quantitative traits under selection without genealogical information. Key words: Fabaceae; heritability; Leguminosae; molecular markers; Prosopis alba; quantitative traits.

Platymiscium is a neotropical legume genus of forest trees in the Pterocarpus clade of the pantropical 'dalbergioid' clade. It comprises 19 species (29 taxa), distributed from Mexico to southern Brazil. This study presents a molecular phylogenetic analysis of Platymiscium and allies inferred from nuclear ribosomal (nrITS) and plastid (trnL, trnL-F and matK) DNA sequence data using parsimony and Bayesian methods. Divergence times are estimated using a Bayesian method assuming a relaxed molecular clock (multidivtime). Within the Pterocarpus clade, new sister relationships are recovered: Pterocarpus + Etaballia, Inocarpus + Tipuana and Paramachaerium + Maraniona. Our results support monophyly of Platymiscium, which is resolved into three major clades, each with distinct geographic ranges and ecological preferences. Diversification in Platymiscium has been driven by habitat fragmentation, invasion of novel geographic regions, and ecological diversification, revealing general patterns of diversification in the neotropics. We hypothesize that Platymiscium arose in dry habitats of South America and radiated northward. The Amazon basin was invaded twice both within the last 5.6 My and Central America twice before the closure of the Isthmus of Panama. Divergence times of the P. pubescens complex, restricted to seasonally dry tropical forests of South America, support pre-Pleistocene divergence in this biome. Key words: biogeography; divergence times; Isthmus of Panama; Leguminosae; neotropics; phylogenetics; Platymiscium; Pterocarpus clade; seasonally dry tropical forests.

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.sub.ST]), whereas S. genistoides subsp. genistoides had a low FST. 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. Key words: chloroplast microsatellites; cpSSRs; Leguminosae; Messinian salinity crisis; northern Morocco; phylogeography; southern Spain; Stauracanthus.

Genetic variation in 27 populations of Ulex species from southern Spain and northern Morocco (Betic-Rif arc) was assessed using 11 chloroplast microsatellite (cpSSR) markers, which revealed 47 different haplotypes. These nonrecombinant, haploid markers allow measurement of genetic variation in closely related species of Ulex where molecular phylogenetic analyses have not provided a clear view of interspecific relationships. Discriminant analysis indicates that the haplotypes are useful to differentiate among species, and analysis of molecular variance (AMOVA) shows high levels of differentiation among populations. The minimum spanning tree (MST), that represents the connections between the haplotypes, suggests that the eastern Rifean U. africanus haplotypes are more genetically related than those from southern Spain. The latter may have lost genetic diversity while colonizing new habitats, eventually differentiating entiating into U. baeticus and U. scaber. Hybridization between these populations, followed by polyploidization, may have originated the tetraploids (U. congestus and U. borgiae) that colonized new habitats associated with acidic rocks. Separate groupings of U. scaber populations may indicate multiple origins from different stocks. Diversification in this group of Ulex species could be related to the opening of the Alboran Sea by Middle Miocene, when the populations from Morocco and Spain became isolated ,torn each other. Key words: chloroplast microsatellitcs; cpSSRs; genetic variation; Leguminosae; northern Morocco; southern Spain; Ulex.

For an introduced plant species to become invasive, it must be able to reproduce even in initially small populations. We tested for Allee effects (reduced reproductive performance of individuals in small populations) in the nonclonal, buzz-pollinated shrub Senna didymobotrya in its invasive range in South Africa. The species is self-compatible, but we found that in its invasive range in South Africa it requires pollinators to set seed. Nearly all stigmas (90%) received pollen, but natural fruit set was very low (3-20%). Pollen receipt and fruit set were not significantly correlated with population size. We thus found no evidence for an ecological Allee effect arising from pollen limitation in small populations. Offspring seedling performance, measured in terms of stem volume and leaf area, was also not significantly correlated with the number of plants in the source population, indicating that genetic Allee effects, such as inbreeding depression, are either absent or of such a small magnitude that they would be unlikely to limit further spread of S. didymobotrya in South Africa. Key words: alien plant; Baker's rule; Cassia didymobotrya; nonindigenous plant; pollination; population size; Xylocopa spp.

The 5S and the 18S-25S rRNA genes were physically mapped by fluorescent in situ hybridization (FISH) in all botanical varieties of cultivated peanut Arachis hypogaea (2n = 4x = 40), in the wild tetraploid A. monticola, and in seven wild diploid species considered as putative ancestors of the tetraploids. A detailed karyotype analysis including the FISH signals and the heterochromatic bands was carried out. Molecular cytogenetic landmarks are provided for the construction of a FISH-based karyotype in Arachis species. The size, number, and chromosome position of FISH signals and heterochromatic bands are similar in all A. hypogaea varieties and A. monticola, but vary among the diploid species. Genome constitution of the species is discussed and several chromosome homeologies are established. The bulk of the chromosome markers mapped, together with data on geographical distribution of the taxa, suggest that peanut originated upon domestication of A. monticola and evidence that the diploids A. duranensis and A, ipaensis are the most probable ancestors of both tetraploid species. Allopolyploidy could have arisen by a single event or, if by multiple events, always from the same diploid species. Key words: Arachis; FISH mapping: heterochromatin: karyotypes: Leguminosae; peanut origin: rRNA genes.

Acacia aroma and A. macracantha are closely related species that inhabit northern and central Argentina. The reproductive barriers between them seem to be weak. They exhibit low genetic differentiation, high levels of interspecific gene flow, and extensive areas of sympatry. Isoenzymatic approaches were used to evaluate the population structure and mating system parameters in natural Argentine populations of A. aroma and A. macracantha and to provide new tools for the analysis of relationships between these two species. All studied populations had high levels of genetic variability and no significant departures from Hardy-Weinberg expectations, but the two species did not differ from each other. Most variability occured within populations. Mating system analysis showed high levels of outcrossing, no biparental inbreeding, and a high probability that individuals within progeny arrays are full rather than half sibs. In all A. aroma and A. macracantha populations, polymorphic loci had the same allelic variants, and no geographic or genetic isolation between species was found. The results favor the hypothesis that these two entities represent a single polymorphie species rather than two distinct species. Key words: Acacia; Argentina; Fabaceae; genetic variability; isozymes; mating system parameters; population structure.

The anomalous systematic position of Swartzieae at the base of Papilionoideae is correlated with unusual developmental features: one petal or none; a ring meristem; polystemony; heterostemony; little or no alignment of stamens as antesepalous or antepetalous; multicarpely; and absence of unidirectional order of organs except in the calyx. Symmetry is zygomorphic throughout development. Floral ontogeny of four species of Swartzia reveals five sepals are initiated successively, beginning abaxially, but intercalary growth below the separate sepals forms a tubular calyx that splits irregularly, a feature typifying the genus. A single petal is initiated adaxially or may be missing altogether (in S. sericea). The apex enlarges and forms a ring meristem concurrently with carpel initiation. Several large-stamen primordia (2-15, according to the species) initiate first on the ring, followed by 40-150 small-stamen primordia. The latter initiate in centrifugal order in S. aureosericea and S. laurifolia or in acropetal order in S. sericea and S. madagascariensis. While ring meristems are considered to be homologous among Neotropical species studied as well as in the African S. madagascariensis, they vary in extent, duration, order of initiation, and productivity. Swartzieae is unlikely to be ancestral to the rest of Papilionoideae, based on radically differing floral ontogeny in the two groups. Key words: Bobgunnia; development; Fabaceae; flower; initiation; Leguminosae; Papilionoideae; ring meristem; ontogeny; polystemony; Swartzia; Swartzieae.

The karyotypes of 10 species and one variety of South American Lathyrus were determined and compared with those obtained of five entities from the Northern Hemisphere. Although all the species have a chromosome number of 2n = 14, they could be differentiated by their karyotype formula and quantitative parameters of the karyotypes. Phenetic distance and principal component analysis showed that in spite of the differences observed among entities, they can be grouped in clusters that coincide with the taxonomic sections established by F. K. Kupicha and with the life cycle of the species. South American species form a homogeneous group and can be distinguished by the presence of a subtelocentric pair, which has a macrosatellite in the long arm, and the lack of a short metacentric pair characteristic of most species of the Northern Hemisphere. From an evolutionary point of view, variation in total chromosome length without major changes in the karyotype formula suggests that changes in the amounts of genomic DNA are proportional to the relative length of each chromosome arm and that species of Notolathyrus evolved in a concerted fashion. Variation in genome size, however, is congruent with morphological variation of some reproductive organs as well as with the life cycle and minimum generation time, as predicted by the nucleotype hypothesis. Key words: concerted evolution; karyotypes; Lathyrus; Leguminosae; Notolathyrus; nucleotypes; South America.

To set up an in situ conservation strategy for Phaseolus lunatus, we analyzed the genetic structure of 29 populations in the Central Valley of Costa Rica. Using 22 enzyme loci, we quantified the proportion of polymorphic loci ([P.sub.p]), the mean number of alleles per locus (A), and the mean effective number of alleles per locus ([A.sub.o]), which equaled to 10.32%, 1.10, and 1.05, respectively. The total heterozygosity ([H.sub.T), the intrapopulation genetic diversity ([H.sub.s]), and the interpopulation genetic diversity ([D.sub.ST) were 0.193, 0.082, and 0.111, respectively. The genotypic composition of the analyzed populations showed a deviation from the Hardy-Weinberg proportions ([F.sub.IT] = 0.932). This disequilibrium was due to either genetic differentiation between populations ([F.sub.ST] = 0.497) or nonrandom mating within populations ([F.sub.IS] = 0.866). From the level of genetic differentiation between populations and the private alleles frequencies estimates, gene flow was calculated: [Nm.sub.w] = 0.398 and [Nm.sub.s] = 0.023, respectively. The results suggested that wild Lima bean maintains most of its isozyme variation among populations. Significant positive correlation was observed between population size and [P.sub.p], A, and [H.sub.o] (observed heterozygosity), whereas no correlation was observed with the average fixation index of population (F). The loss of genetic variability in populations was attributed to inbreeding and the bottleneck effects that characterized the target populations. In situ conservation and management procedures for wild Lima bean arc discussed. Key words: Costa Rica; Fabaceae; gene flow; genetic diversity; in situ conservation; isozymes; Lima bean; Phaseolus lunatus; population size.

We used nucleotide sequences from the internal transcribed spacers and 5.85 gene of nuclear ribosomal DNA to test competing phylogenetic and biogeographic hypotheses in Gleditsia. Eleven of 13 Gleditsia species were sampled, along with two species of its sister genus, Gymnocladus. Analyses of ITS data and of a combined data set that included sequences of ITS and two chloroplast genes supported several conclusions that were interpreted in light of fossil data and current legume phylogeny. Gleditsia and Gymnocladus appear to have originated in eastern Asia during the Eocene. Eastern North American species of both genera most likely evolved from ancestors that migrated across the Bering land bridge, but the eastern Asian/eastern North American disjunction appears to be much older in Gymnocladus than in Gleditsia. Gleditsia amorphoides, from temperate South America, is sister to the rest of the genus, suggesting early long-distance dispersal from Asia. The remainder of Gleditsia is divided into three unresolved clades, possibly indicating a split early in the evolution of the genus. Two of those clades contain only Asian species, and one contains Asian and North American species. The North American species, Gleditsia triacanthos and Gleditsia aquatica, are polymorphic and paraphyletic with respect to their ITS and cpDNA sequences, which suggests recent diversification. Key words: biogeography; Gleditsia; Gymnocladus; internal transcribed spacer; Leguminosae; molecular phylogeny; ribosomal DNA.

Leucaenas are fast-growing neotropical trees and shrubs used as forage and fuelwood. This study tested the seed cross- compatibilities among 15 of the 16 well-defined species of the paleopolyploid genus Leucaena Bentham (Leguminosae: Mimosoideae) to determine if genes of interest could be transferred between species. Twelve diploid and four tetraploid taxa were artificially hybridized in 118 of all 120 possible two-way combinations and 31 of 32 possible self- and intraspecific matings, with 58,218 floret pollinations. Ninety-one combinations (77%) produced well-filled seed, and 73 of these combinations were grown and validated in the field. The qualitative crossability within ploidy levels (diploid, triploid, tetraploid) was 78%, 75%, and 100%, respectively, whereas quantitative crossability (relative to intraspecific matings) of successful matings within identical ploidy levels was 32%, 11%, and 73%, respectively. One-third (34%) of the 114 interspecific combinations tested in both directions exhibited weak unilateral incompatibility (UI), due, in part, to reciprocal differences in stylar length. Other mechanisms for UI (embryo-endosperm imbalance, sex-specific use of self-compatible parents) were ruled out. Wide hybridization appears to be 'buffered' by polyploidy in the genus.