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Fig.. Papilionoideae I. A, Castanospermum australe; B, Petaladenium urceoliferum; C, Pterodon abruptus; D, Swartzia acutifolia; E, Trischidium molle; F, Exostyles venusta; G, Harleyodendron unifoliolatum; H, Haplormosia monophylla; I, Ormosia lewisii; J, Harpalyce lanata; K, Leptolobium brachystachyum; L, Camoensia brevicalyx.-Photos: A-G & I-K, Domingos Cardoso; H, Jan Wieringa; L, André van Proosdij. 

Fig.. Papilionoideae I. A, Castanospermum australe; B, Petaladenium urceoliferum; C, Pterodon abruptus; D, Swartzia acutifolia; E, Trischidium molle; F, Exostyles venusta; G, Harleyodendron unifoliolatum; H, Haplormosia monophylla; I, Ormosia lewisii; J, Harpalyce lanata; K, Leptolobium brachystachyum; L, Camoensia brevicalyx.-Photos: A-G & I-K, Domingos Cardoso; H, Jan Wieringa; L, André van Proosdij. 

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The classification of the legume family proposed here addresses the long-known non-monophyly of the traditionally recognised subfamily Caesalpinioideae, by recognising six robustly supported monophyletic subfamilies. This new classification uses as its framework the most comprehensive phylogenetic analyses of legumes to date, based on plastid matK...

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... The Fabaceae family (legumes) is a large group of flowering plants, with around 19,500 species. This family is remarkably diverse and stands out as the second most economically important family of crop plants after grasses (Azani et al. 2017;Thorne 2002). The notable grain legumes -such as chickpea (Cicer arietinum), pea (Pisum sativum), peanut (Arachis hypogaea), common bean (Phaseolus vulgaris), and soybean (Glycine max) -play a critical role in human and animal nutrition, as well as in industrial applications. ...
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Soybean (Glycine max) and common bean (Phaseolus vulgaris) diverged approximately 19 million years ago. While these species share a whole-genome duplication (WGD), the Glycine lineage experienced a second, independent WGD. Despite the significance of these WGDs, their impact on gene families related to oil-traits remains poorly understood. Here, we report an in-depth investigation of oil-related gene families in soybean, common bean, and twenty-eight other legume species. We adopted a systematic approach that included transcriptome and co-expression analysis, identification of orthologous groups, gene duplication modes and evolutionary rates, and family expansions and contractions. We curated a list of oil candidate genes and found that 91.5% of the families containing these genes expanded in soybean in comparison to common bean. Notably, we observed an expansion of triacylglycerol (TAG) biosynthesis (~3:1) and an erosion of TAG degradation (~1.4:1) families in soybean in comparison to common bean. In addition, TAG degradation genes were two-fold more expressed in common bean than in soybean, suggesting that oil degradation is also important for the sharply contrasting seed oil contents in these species. We found 17 transcription factor hub genes that are likely regulators of lipid metabolism. Finally, we inferred expanded and contracted families and correlated these patterns with oil content found in different legume species. In summary, our results do not only shed light on the evolution of oil metabolism genes in soybean, but also present multifactorial evidence supporting the prioritization of candidates for crop improvement.
... Perez-Escobar et al. (2023) presented a novel hypothesis of biogeography of orchid plants, proving that taxon sampling depth and outgroups influence results. There is thus an urgent need for global collaboration to undertake large-scale phylogenomic work in other plant lineages, as in the case of Fabaceae (Azani et al., 2017), Lamiaceae (Boachon et al., 2018), and Poaceae (Soreng et al., 2017(Soreng et al., , 2022; and Carex (Roalson et J o u r n a l P r e -p r o o f al., 2021). Achieving large-scale sampling efforts are vital to enhance scientific comprehension of lineage diversification with a large number of species, leveraging existing genomic data and increasing the scope of cooperation in research (Givnish et al., 2016;Thompson et al., 2023). ...
... Furthermore, the distribution of Chamaecrista is unique in being the only nodulated caesalpinioid genus which has species that have colonized temperate regions (Sprent et al., 2013). All Chamaecrista species so far examined form symbiotic root nodules with nitrogen-fixing bacteria collectively known as rhizobia (Gyaneshwar et al., 2011;Peix et al., 2015;Sprent et al., 2017), while related genera comprising the Cassia clade (Azani et al., 2017), p. ex. Cassia L., and Senna Mill. ...
... In the present study, the tree species C. eitenorum, which is native to SDTFs, had the apparently unique property of being nodulated by strains of Paraburkholderia related to those that nodulate Mimosa and Calliandra spp. in the same environment (Bontemps et al., 2010;Silva et al., 2018); indeed, the fact that these isolates were also capable of nodulating C. duartei and C. ensiformis in crossinoculation studies suggests that Chamaecrista-Paraburkholderia symbioses might be relatively abundant, at least among the tree species. Further to this, the C. eitenorum-Paraburkholderia interaction is the first confirmed report of a beta-rhizobial symbiosis in nodules from a Caesalpinioideae species outside the Mimosoid clade (Azani et al., 2017), and it is also the first report of FTs being occupied by this type of microsymbiont, which demonstrates that the FT phenotype is not controlled by any particular rhizobial type (e.g. Bradyrhizobium), but is an inherent plant characteristic, which might be expected if the 'host controls the party' (Ferguson et al., 2019). ...
Article
All non-mimosoid nodulated genera in the legume subfamily Caesalpinioideae confine their rhizobial symbionts within cell wall-bound “fixation threads” (FTs). The exception is the large genus Chamaecrista in which shrubs and subshrubs house their rhizobial bacteroids more intimately within symbiosomes, whereas large trees have FTs. This study aimed to unravel the evolutionary relationships between Chamaecrista growth habit, habitat, nodule bacteroid type, and rhizobial genotype. The growth habit, bacteroid anatomy, and rhizobial symbionts of 30 nodulated Chamaecrista species native to different biomes in the Brazilian state of Bahia, a major centre of diversity for the genus, was plotted onto an ITS-TrnL-F-derived phylogeny of Chamaecrista. The bacteroids from most of the Chamaecrista species examined were enclosed in symbiosomes (SYM-type nodules), but those in arborescent species in the section Apoucouita, at the base of the genus, were enclosed in cell wall material containing homogalacturonan (HG) and cellulose (FT-type nodules). Most symbionts were Bradyrhizobium genotypes grouped according to the growth habits of their hosts, but the tree, C. eitenorum, was nodulated by Paraburkholderia. Chamaecrista has a range of growth habits that allow it to occupy several different biomes and to co-evolve with a wide range of (mainly) bradyrhizobial symbionts. FTs represent a less intimate symbiosis linked with nodulation losses, so the evolution of SYM-type nodules by most Chamaecrista species may have (a) aided the genus-wide retention of nodulation, and (b) assisted in its rapid speciation and radiation out of the rainforest into more diverse and challenging habitats.
... It is well known that numerous rhizobial genes contribute to plant nodulation (nod) and nitrogen fixation (nif, fix) processes, and some of them are used for characterization of symbiotic relationships and host range prediction 1,14,21,53,[67][68][69][70][71][72][73][74][75] . Among these genes, nodC and nifH are used in most studies to classify rhizobial strains to particular symbiotic variants (sv.). ...
... In our earlier study, we classified these clover root isolates to R. leguminosarum species 46 and results obtained in this work allowed us to classify them to the symbiovar trifolii. Generally, phylogenetic analysis of two symbiotic nodC and nifH genes are sufficient to assign strains to a particular symbiovar, as it was confirmed for many isolates from different species 1,14,21,53,[67][68][69][70][71][72][73][74][75] . For an example based on the nodC sequences, isolates from Phaseolus vulgaris were classified to symbiovars phaseoli and gallicum 70 . ...
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Red clover (Trifolium pratense L.) is a forage legume cultivated worldwide. This plant is capable of establishing a nitrogen-fixing symbiosis with Rhizobium leguminosarum symbiovar trifolii strains. To date, no comparative analysis of the symbiotic properties and heterogeneity of T. pratense microsymbionts derived from two distinct geographic regions has been performed. In this study, the symbiotic properties of strains originating from the subpolar and temperate climate zones in a wide range of temperatures (10–25 °C) have been characterized. Our results indicate that all the studied T. pratense microsymbionts from two geographic regions were highly efficient in host plant nodulation and nitrogen fixation in a wide range of temperatures. However, some differences between the populations and between the strains within the individual population examined were observed. Based on the nodC and nifH sequences, the symbiotic diversity of the strains was estimated. In general, 13 alleles for nodC and for nifH were identified. Moreover, 21 and 61 polymorphic sites in the nodC and nifH sequences were found, respectively, indicating that the latter gene shows higher heterogeneity than the former one. Among the nodC and nifH alleles, three genotypes (I–III) were the most frequent, whereas the other alleles (IV–XIII) proved to be unique for the individual strains. Based on the nodC and nifH allele types, 20 nodC-nifH genotypes were identified. Among them, the most frequent were three genotypes marked as A (6 strains), B (5 strains), and C (3 strains). Type A was exclusively found in the temperate strains, whereas types B and C were identified in the subpolar strains. The remaining 17 genotypes were found in single strains. In conclusion, our data indicate that R. leguminosarum sv. trifolii strains derived from two climatic zones show a high diversity with respect to the symbiotic efficiency and heterogeneity. However, some of the R. leguminosarum sv. trifolii strains exhibit very good symbiotic potential in the wide range of the temperatures tested; hence, they may be used in the future for improvement of legume crop production.
... In the case of R. pseudoacacia, it is possible that the high levels of free feeding damage are caused by native insects specialized for feeding on the Fabaceae. The Fabaceae is the world's third largest land plant family, and its species are native to every temperate world region (Azani et al. 2017). Therefore, there is considerable potential for native herbivore species to extend their host range to include introduced R. pseudoacacia. ...
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There is a long history of humans either intentionally or accidentally moving plant species to areas outside of their native ranges. In novel environments, populations of many of these plant species exhibit explosive population growth and spread, in part due to the absence of coevolved enemies such as herbivorous insects. However, over time such enemies can ‘catch up' with their host and re‐establish host–herbivore relationships. Though this phenomenon has been documented in several systems, little evidence exists on how this re‐assembly of enemies results in increased levels of herbivory. In this study we focus on the case of black locust Robinia pseudoacacia, a sparsely populated tree species when growing on undisturbed sites in its limited native range in the eastern USA but a highly invasive species, especially in disturbed environments, in most temperate world regions. We recorded folivore damage on invasive populations in five continents, including both native and invaded portions of North America. Here, we investigated 1) how total foliage damage and damage caused by different groups of folivores differs among regions; 2) how seasonal development of folivore damage differs among regions; 3) how folivory varies with distance from the native range within North America; and 4) how the number of recorded specialist folivores correlates with the amount of folivory. We observed strong differences among regions in the amount and type of folivore damage, with the native range experiencing the highest damage, especially that caused by the native chrysomelid beetle Odontota dorsalis, which is limited to the native and invaded North American range of R. pseudoacacia. Among world regions, total folivory is negatively associated with the distance from the native range and positively associated with the number of established R. pseudoacacia specialist folivore species, supporting the hypothesis that global patterns of herbivore invasions are associated with diminished enemy release.
... Legumes (Fabaceae), the third largest family of seed plants, typically have compound leaves 19 . The family is currently divided into six subfamilies: Cercidoideae, Detarioideae, Duparquetioideae, Dialioideae, Caesalpinioideae and Papilionoideae, with most having pinnate compound leaves in a variety of forms (except for Cercidoideae) (Fig. 1a) 20 . For instance, unipinnate leaves can be either paripinnate (when ending in two leaflets) or imparipinnate (when ending in one leaflet) with alternate or opposite leaflet arrangements (Fig. 1a). ...
Article
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Plant lateral organs are often elaborated through repetitive formation of developmental units, which progress robustly in predetermined patterns along their axes. Leaflets in compound leaves provide an example of such units that are generated sequentially along the longitudinal axis, in species-specific patterns. In this context, we explored the molecular mechanisms underlying an acropetal mode of leaflet initiation in chickpea pinnate compound leaf patterning. By analyzing naturally occurring mutants multi-pinnate leaf1 (mpl1) that develop higher-ordered pinnate leaves with more than forty leaflets, we show that MPL1 encoding a C2H2-zinc finger protein sculpts a morphogenetic gradient along the proximodistal axis of the early leaf primordium, thereby conferring the acropetal leaflet formation. This is achieved by defining the spatiotemporal expression pattern of CaLEAFY, a key regulator of leaflet initiation, and also perhaps by modulating the auxin signaling pathway. Our work provides novel molecular insights into the sequential progression of leaflet formation.
... The region's geomorphological heterogeneity and palaeoclimatic changes have contributed to its unique biodiversity (Prado 2003, Queiroz 2006, Pennington et al. 2009, Guedes et al. 2014, Oliveira et al. 2015, Werneck et al. 2015, Azani et al. 2017, Queiroz et al. 2017, Fernandes et al. 2022. Diverse hypotheses and factors have been suggested to explain the diversification, demography, genetic structuring, and distribution of genetic diversity of species and populations within the Caatinga. ...
Article
The Caatinga biogeographical region, located in the semi-arid region of northeastern Brazil, is characterized by high levels of cactus diversity and endemism. In this study, we investigated the genetic diversity, differentiation, and phylogeography of three Tacinga species (Tacinga inamoena, Tacinga subcylindrica, and Tacinga palmadora), which are endemic and widely distributed in the Caatinga. We collected 222 samples from 108 locations and used plastid (3ʹrps16-5ʹtrnK, PsbE-PetL) and nuclear (PhyC) molecular markers for a multilocus approach. Our results suggest that diversification of the three Tacinga species occurred during the middle Pleistocene, with the Southern Sertaneja Depression ecoregion of the Caatinga being identified as the ancestral area. Interestingly, analyses of genetic structure did not show any evidence that the major river in the region (the Sao Francisco River) and the Caatinga ecoregions acted as biogeographical barriers for gene flow in these species. This suggests that the three Tacinga species exhibited different responses to widely accepted geographical and geomorphological barriers known to affect other taxonomic groups in the Caatinga. Overall, our findings provide new insights into the biogeographical history of the Caatinga and the factors that have shaped diversification and genetic structure of species in this region. Our study underlines the importance of considering the unique natural history and biological characteristics of individual taxonomic groups when investigating the biogeography of the Caatinga.
... Ecologically successful taxa evolve successful and efficient N assimilation systems to survive in severe habitats or compete for nourishment. Approximately 90% of the species within the family Leguminosae can fix atmospheric N through a symbiotic association with soil bacteria and have become widespread through the most spectacular radiations 19 . Orchids are one of the very few flowering plant lineages that have been able to successfully colonize epiphytic or lithophytic niches, clinging to trees or rocks and growing in dry conditions using crassulacean acid metabolism 20 . ...
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The Asteraceae (daisy family) is one of the largest families of plants. The genetic basis for its high biodiversity and excellent adaptability has not been elucidated. Here, we compare the genomes of 29 terrestrial plant species, including two de novo chromosome-scale genome assemblies for stem lettuce, a member of Asteraceae, and Scaevola taccada, a member of Goodeniaceae that is one of the closest outgroups of Asteraceae. We show that Asteraceae originated ~80 million years ago and experienced repeated paleopolyploidization. PII, the universal regulator of nitrogen-carbon (N-C) assimilation present in almost all domains of life, has conspicuously lost across Asteraceae. Meanwhile, Asteraceae has stepwise upgraded the N-C balance system via paleopolyploidization and tandem duplications of key metabolic genes, resulting in enhanced nitrogen uptake and fatty acid biosynthesis. In addition to suggesting a molecular basis for their ecological success, the unique N-C balance system reported for Asteraceae offers a potential crop improvement strategy.
... Asteraceae reaches 20% between ad 200 and 700, a period that corresponds to higher cultural and agricultural development in Oxpemul, determined archaeologically (Folan et al. 2015). The subfamily Caesalpinioideae (Azani et al. 2017) is present with a low percentage (< 5%), an abundant taxa in secondary vegetation and lowland forests. As part of the non-pollen palynomorphs (NPP), an abundance of fungal spores was identified reaching values above 20%, like other paleoenvironmental records of the YP. ...
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Multiple approaches were used to investigate agricultural and forestry practices of the Preclassic Maya in the Petén Campechano, southwestern Yucatan Peninsula, Mexico. Palaeoenvironmental inferences were based on pollen and geochemical data from sediment cores collected in lakes Silvituc and Uxul, and the Oxpemul Reservoir. These water bodies are near three archaeological sites that supported agricultural activity between ca. 900 bc and ad 750. After ca. ad 500 the area was under the control of the Kaan Dynasty. These sites show patterns similar to those in sediments from Lake Petén Itzá, northern Guatemala, but they are different from records from the northeastern part of the Peninsula at Lake Chichancanab. Changes in the patterns of abundance, increase and decrease of maize pollen, relative to that of other crops (Cucurbitaceae and Chenopodiaceae) reflect changes with respect to their cultivation and importance. High clay and gypsum content in the sediments may be related to deforestation, agriculture, and intervals of greater rainfall. Changes in sediment elemental concentrations were associated with droughts and human activities during the Classic Period after ad 250. The Classic droughts were more severe and prolonged than those of the Preclassic and thus had a greater impact for inhabitants of the Petén Campechano. This led to a cultural collapse by the seventh century, whereas settlements farther north on the Yucatan Peninsula persisted until the tenth century. Gradually increasing precipitation during the Postclassic (ad 1350–1850), contemporary with the Little Ice Age (LIA), is consistent with the paleoclimate history of the Guatemalan Petén. The response of ecosystems to droughts revealed the water vulnerability of the region and its influence on the sustainability of Maya settlements.
... To investigate the extent of conservation of pulvinar slits among legumes, we stained vertical sections of pulvini from various legume species with calcofluor white. The Fabaceae family comprises 2 major subfamilies: the Papilionoideae and the Caesalpinioideae (Azani et al. 2017). Importantly, we detected slit structures in pulvinar CMCs from all 11 species selected and tested from these 2 subfamilies (Supplemental Table S1, Fig. 1H). ...
Article
The cortical motor cells (CMCs) in a legume pulvinus execute the reversible deformation in leaf movement that is driven by changes in turgor pressure. In contrast to the underlying osmotic regulation proper, the cell wall structure of CMCs that contributes to the movement has yet to be characterized in detail. Here, we report that the cell wall of CMCs has circumferential slits with low levels of cellulose deposition, which are widely conserved among legume species. This structure is unique and distinct from that of any other primary cell walls reported so far; thus, we named them "pulvinar slits." Notably, we predominantly detected de-methyl-esterified homogalacturonan inside pulvinar slits, with a low deposition of highly methyl-esterified homogalacturonan, as with cellulose. In addition, Fourier-transform infrared spectroscopy analysis indicated that the cell wall composition of pulvini is different from that of other axial organs, such as petioles or stems. Moreover, monosaccharide analysis showed that pulvini are pectin-rich organs like developing stems and that the amount of galacturonic acid in pulvini is greater than in developing stems. Computer modeling suggested that pulvinar slits facilitate anisotropic extension in the direction perpendicular to the slits in the presence of turgor pressure. When tissue slices of CMCs were transferred to different extracellular osmotic conditions, pulvinar slits altered their opening width, indicating their deformability. In this study, we thus characterized a distinctive cell wall structure of CMCs, adding to knowledge of repetitive and reversible organ deformation as well as the structural diversity and function of the plant cell wall.