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The papilionoid genus Stylosanthes Sw. includes about 50 spp. distributed world wide in the tropics, approximately half of them grow in South America. The present study focuses on South American Stylosanthes. Based on examinations of herbarium specimens, as well as field observations, a total of 25 taxa (23 spp. and 2 varieties) are here described...
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Context 1
... V. The type and shape of the fruit is important to define the species of Stylosanhes. Fruits of some species are illustrated in Fig. ...
Context 2
... Stylosanthes humilis is a native species of Venezuela that has been introduced in many countries as a forage plant, including Brazil, Bolivia and Paraguay. Fig. ...
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Citations
... N-xing forage legumes may increase milk and meat production and even reduce greenhouse gas and enteric methane emissions (Boddey et al. 2020). Some studies even showed a promising response for cultivated Stylosanthes species to increased temperatures due to climate change (Martinez et Several authors have studied group taxonomy (Burkart 1939;Mohlenbrock 1957Mohlenbrock , 1962Ferreira and Costa 1979;'t Mannetje 1984;Costa 2006;Vanni 2017), but these studies often diverge in terms of recognition of some taxa, and the identi cation of species is still challenging. Despite Costa's (2006) During an extensive analysis of the morphology and nomenclature of Stylosanthes species from Brazil (Gissi 2022), it was clear that taxonomic updates were needed. ...
Stylosanthes Sw. is a pantropical genus primarily distributed throughout the Americas, with the Brazilian Cerrado serving as its principal center of diversity. Many of its species hold significant economic potential and are widely utilized as fodder for cattle. Systematic examinations involving both morphology and nomenclature have highlighted the imperative need for taxonomic updates in select species. Given the observed morphological and ecological differences, a taxonomic reevaluation of S. guianensis (Aubl.) Sw. was deemed necessary, leading to the elevation of its varieties to the species level. The taxonomic changes include the new combinations S. microcephala (M.B. Ferreira & Sousa Costa) Gissi, S. pauciflora (M.B. Ferreira & Sousa Costa) Gissi, and S. pubescens (Pilger) Gissi. Furthermore, a neotype for S. nunoi Brandão is designated due to the loss of the holotype. Stylosanthes pohliana Taub. is reinstated as an accepted species, and a novel species within the genus is described. X-ray imaging of leaflets emerges as a valuable tool for taxonomic discrimination. This comprehensive study not only presents nomenclatural modifications but also provides illustrations, comments on diagnostic characteristics, distribution maps, and preliminary IUCN conservation assessments. Additionally, a novel discovery is introduced with the recognition of an elaiosome in the fruits of S. guianensis , constituting a noteworthy addition to the tribe Dalbergieae.
... For this study, specimens of all South American Stylosanthes species deposited in the following herbaria were analyzed: ALCB, ASE, BOTU, CEN, EAC, ESA, HBRA, HUEFS, HVASF, IAN, INPA, L, MFS, NY, P, R, RB, SLUI, SP, UB, US, WAG (acronyms according to Thiers, 2021). The relevant taxonomic literature on Stylosanthes was also consulted (Vogel, 1838;Mohlenbrock, 1958;'t Mannetje, 1984;Costa, 2006;Vanni, 2017;Gissi, 2020). The use of morphological terms mainly follows Harris and Harris (2001) and Radford (1974). ...
A new species of Stylosanthes (Leguminosae – Papilionoideae) from the Chapada das Mesas National Park in Maranhão, Brazil, is described. The new species, S. acicularis, was misidentified as S. angustifolia in the SLUI herbarium. It differs from S. angustifolia especially by inflorescence shape and rostrum length. The new species closely resembles S. hippocampoides by its small capitate inflorescences, absence of a rudimentary axis beneath the flower, only one inner bracteole, loments with only one fertile article and a very short rostrum, and the absence of a marginal vein and possession of up to five secondary veins in the leaflets. However, the two species differ by stem indumentum, bristle length, leaflet shape and venation, and shape of the outer bracteole margin. A morphological description, illustrations—including X-ray imaging of the leaflet venation—and notes on the geographical distribution, habitat, and conservation status of the new species are presented.
... In previous major taxonomic studies of the genus (Mohlenbrock, 1958;Ferreira and Costa, 1979;Vanni, 2017), the indumentum of different Stylosanthes species has been referred to as hirsute, hispid, or pubescent, and sometimes as pilose, setose or hairy-viscous. The structures that make up the indumentum have been referred to as hairs, bristles or tuberculate-bristles. ...
... This system was largely followed by Costa (2006), but with the addition of a fourth variety of S. guianensis and the recognition of another segregate species, S. nunoi. Finally, Vanni (2017) subsumed all the segregate species, except S. hippocampoides, within S. guianensis, within which he recognized only two varieties in South America -S. guianensis var. ...
... Some authors (Mohlenbrock, 1958;Mannetje, 1977Mannetje, , 1984Vanni, 2017) have adopted a broader circumscription of S. guianensis, subsuming within it several other binomials, which in our opinion should be maintained as separate species. Under our concept of the species, S. guianensis is distinguished by the fruit with only one fertile article, a very short rostrum (less than 1 mm), and visible glands at the base (Costa, 2006). ...
Stylosanthes comprises about 50 species, with Brazil being one of the main diversity centers. Stylosanthes guianensis constitutes a problematic taxonomic complex, comprising four varieties of S. guianensis and seven other species. Considering the difficulty of taxon delimitation, this study assessed the potential value of the secretory structures and other anatomical characters to the diagnosis of taxa within the complex. We analyzed the leaflet and stem anatomy and micromorphology of representatives of the S. guianensis complex using light and scanning electron microscopy. Long uniseriate non-glandular hairs occurred on the stems and leaflets of all species. Secretory structures with a dilated base and a tapered distal portion occurred on both stems and leaflets, except in S. guianensis var. canescens. Polysaccharides, lipids and terpenes were histochemically detected in the secretory exudates. In the stem of S. gracilis, chlorenchyma cells were observed to be radially elongated, while in the other taxa studied they were isodiametric. On leaflets of S. guianensis var. pauciflora, S. acuminata, S. aurea, and S. gracilis, the midrib was eccentric and the veins closer to the leaflet margin had larger fiber caps. Anatomical structures proved to be useful for separating taxa. Stylosanthes guianensis var. canescens and S. guianensis var. pauciflora differ from the other varieties of the species and are possibly worthy of recognition at species rank. Our findings also reinforce the status of S. gracilis as a distinct species.
... The genus Stylosanthes Sw. (Leguminosae) belongs to the subfamily Papilionoideae and has a complex systematics, mainly related to the occurrence of natural allopolyploidy (Stace and Edye, 1984;Vanni, 2017). The taxonomy of the genus remains unsettled and controversial, with various authors favoring between 25 and 42 species, with at least 40 additional synonyms (Cameron and Chakraborty, 2004). ...
... Stylosanthes is highly diversified and morphologically polymorphic, having cultivated pantropical species, mostly described for the American continent with two centers of diversification: Mexico and Brazil. Being the latter the main center of origin and diversification for the genus, with more than 30 species, of which 12 are endemic (Stace and Edye, 1984;da Costa and Valls, 2010;Santos-Garcia et al., 2012;Vanni, 2017). Species circumscription and identification are complex in Stylosanthes since many different species have overlapping morphological characters, many of them dubious e/or homoplastic (Costa and Ferreira, 1984;Mannetje, 1984;Vanni, 2017). ...
... Being the latter the main center of origin and diversification for the genus, with more than 30 species, of which 12 are endemic (Stace and Edye, 1984;da Costa and Valls, 2010;Santos-Garcia et al., 2012;Vanni, 2017). Species circumscription and identification are complex in Stylosanthes since many different species have overlapping morphological characters, many of them dubious e/or homoplastic (Costa and Ferreira, 1984;Mannetje, 1984;Vanni, 2017). This makes it necessary to use additional data to taxonomy, such as molecular markers (Liu et al., 1999;Liu and Musial, 2001), molecular phylogeny (Vander Stappen et al., 1999;Vander Stappen et al., 2002), genomics or cytogenetics (Marques et al., 2018;Franco et al., 2020). ...
Allopolyploidy is widely present across plant lineages. Though estimating the correct phylogenetic relationships and origin of allopolyploids may sometimes become a hard task. In the genus Stylosanthes Sw. (Leguminosae), an important legume crop, allopolyploidy is a key speciation force. This makes difficult adequate species recognition and breeding efforts on the genus. Based on comparative analysis of nine high-throughput sequencing (HTS) samples, including three allopolyploids (S. capitata Vogel cv. “Campo Grande,” S. capitata “RS024” and S. scabra Vogel) and six diploids (S. hamata Taub, S. viscosa (L.) Sw., S. macrocephala M. B. Ferreira and Sousa Costa, S. guianensis (Aubl.) Sw., S. pilosa M. B. Ferreira and Sousa Costa and S. seabrana B. L. Maass & 't Mannetje) we provide a working pipeline to identify organelle and nuclear genome signatures that allowed us to trace the origin and parental genome recognition of allopolyploids. First, organelle genomes were de novo assembled and used to identify maternal genome donors by alignment-based phylogenies and synteny analysis. Second, nuclear-derived reads were subjected to repetitive DNA identification with RepeatExplorer2. Identified repeats were compared based on abundance and presence on diploids in relation to allopolyploids by comparative repeat analysis. Third, reads were extracted and grouped based on the following groups: chloroplast, mitochondrial, satellite DNA, ribosomal DNA, repeat clustered- and total genomic reads. These sets of reads were then subjected to alignment and assembly free phylogenetic analyses and were compared to classical alignment-based phylogenetic methods. Comparative analysis of shared and unique satellite repeats also allowed the tracing of allopolyploid origin in Stylosanthes, especially those with high abundance such as the StyloSat1 in the Scabra complex. This satellite was in situ mapped in the proximal region of the chromosomes and made it possible to identify its previously proposed parents. Hence, with simple genome skimming data we were able to provide evidence for the recognition of parental genomes and understand genome evolution of two Stylosanthes allopolyploids.
... The 15 species sampled that are not endemic to Brazil also occur in Bolivia, Colombia, the Guianas, Ecuador, Peru, and Venezuela. The plant specimens were identified using the keys of Costa (2006), Costa et al. (2008), andVanni (2017). Table 1 presents a list of the taxa examined and information from their specimen labels. ...
... The genus Stylosanthes (Fabaceae) includes ∼48 species (Costa & Ferreira, 1984;Mohlenbrock, 1957;Mannetje, 1984) and is originally from Central and South America, being distributed from Mexico to Argentina (Stace & Edye 1984). Recently, a total of 23 species were described for South America (Vanni, 2017), with 11 of them being diploid (2n = 2x = 20), eight being tetraploid (2n = 4x = 40), and the remaining species being unknown (Maass & Sawkins, 2004). The reproduction of Stylosanthes preferably occurs by self-pollination (Bray & Hutton, 1976), with few records of low cross-pollination rates from 2 to 6% (Miles, 1985;Stace, 1984) to 31% for some species (Santos-Garcia, Resende, Chiari, Zucchi, & de Souza, 2011). ...
... In Argentina, eight Stylosanthes species grow naturally (Burkart, 1943;Vanni, 2017), and at least three of them are forage species: S. guianensis (Aubl) Sw., S. hippocampoides Mohlenbr., and S. scabra Vogel (Maass & Sawkins, 2004;Huang, Liu, & Changjun, 2017). The distribution of the first two species covers the northeastern region of Argentina (NEA) (Vanni, 2017). ...
... In Argentina, eight Stylosanthes species grow naturally (Burkart, 1943;Vanni, 2017), and at least three of them are forage species: S. guianensis (Aubl) Sw., S. hippocampoides Mohlenbr., and S. scabra Vogel (Maass & Sawkins, 2004;Huang, Liu, & Changjun, 2017). The distribution of the first two species covers the northeastern region of Argentina (NEA) (Vanni, 2017). This is a subtropical region where three phytogeographic provinces-Chaqueña, Espinal, and Paranaense-converge (Cabrera, 1976), and it contains several ecoregions (Burkart, Bárbaro, Sánchez, & Gómez, 1999), giving rise to wide ecotones and resulting in very diverse patterns of floristic variability (Burkart, 1943;Carnevali, 1994). ...
The Stylosanthes genus is native to Central and South America and contains several of the most important tropical legumes used for forage. The southernmost natural distribution of the genus includes the northeast of Argentina, a subtropical region where several phytogeographic provinces and ecoregions converge, giving rise to particular environmental conditions. Here, we used inter‐simple sequence repeat markers to evaluate the genetic diversity and genetic structure of four and eight natural populations from northeast of Argentina of Stylosanthes guianensis (Aubl.) Sw. and S. hippocampoides Mohlenbr., respectively. We also modeled the potential ecological niches to obtain environmental information regarding new collection areas and future crop areas. High genetic diversity and specific bands were found for both species. The summary statistics were relatively higher for S. guianensis than for S. hippocampoides. In both species, more genetic diversity was present among populations than within them, and correlation between genetic and geographic distances was observed. Populations were grouped according to their respective species, and their geographical source. The results suggest that collecting a few individuals from a large number of different sites would be an appropriate method for conserving the diversity of these species, with emphasis on those areas with the highest levels of genetic diversity. The modeling of ecological niches suggested mean temperature of the coldest quarter as the most important bioclimatic variable influencing S. guianensis distribution, and temperature seasonality as the variable influencing S. hippocampoides distribution, and showed new potential areas for future collections and as potential crop areas of the species.
... Papilionoideae Cologania [80] Papilionoideae Condylostylis [48] Papilionoideae Coursetia [80] Papilionoideae Crotalaria [80,112] Papilionoideae Cyclolobium [116,119] Papilionoideae Dalbergia [80,132] Papilionoideae Dalea [113] Papilionoideae Desmodium [97,99,100] Papilionoideae Dioclea [80] Papilionoideae Discolobium [80, 102. 109] Papilionoideae Dolichopsis [123] Papilionoideae Eriosema [105] Papilionoideae Erythrina [120] Papilionoideae Galactia [111] Papilionoideae Geoffroea [80] Papilionoideae Helicotropis [48] Papilionoideae Holocalyx [80] Papilionoideae Indigofera [114] Papilionoideae Lathyrus [80] Papilionoideae Lonchocarpus [114,118] Papilionoideae Luetzelburgia [134] Papilionoideae Lupinus [80] Papilionoideae Machaerium [80] Papilionoideae Macroptilium [98] Papilionoideae Medicago [80] Papilionoideae Melilotus [80] Papilionoideae Myrocarpus [80,115] Papilionoideae Nissolia [80] Papilionoideae Muellera [80,114,118] Papilionoideae Otholobium [80] Papilionoideae Phaseolus [123] Papilionoideae Poisonia [80] Papilionoideae Poiretia [80] Papilionoideae Pterocarpus [80] Papilionoideae Rhynchosia [95] Papilionoideae Sesbania [114] Papilionoideae Stylosanthes [104,108] Papilionoideae Sweetia [80] Papilionoideae Tephrosia [80] Papilionoideae Trifolium [80] Papilionoideae Vicia [80,110] Papilionoideae Vigna [48,123]; Delgado-Salinas et al. 2011 ...
... Most of the Papilionoideae are exclusive of the Humid Chaco and their most diverse genera occur in temporarily inundated soils, in river coasts, and savannas or grasslands, such as Aeschynomene, Centrosema, Chaetocalyx DC., Desmodium, Discolobium, Eriosema, Galactia, Stylosanthes Sw., Zornia J.F. Gmel., and Vicia L. [96,97,101,102,103,104,105,106,107,108,109,110,111]. The genus Crotalaria comprises several generalist entities in Chaco, but it is remarkable the existence of a recently renamed endemic species in Sierra Chaco [112]. ...
... Some genera of Papilionoideae, either monotypic or poorly represented in the Gran Chaco, such as Myrocarpus Allemão, Amburana Schwacke & Taub. (a SDTF indicator species [18]), Machaerium, Sesbania, Nissolia Jacq., Discolobium Benth., Lonchocarpus Kunth, Muellera L.f., Pterocarpus Jacq., Cyclolobium Benth., Dalbergia L. f., Erythrina L., and Sweetia Spreng., are mainly confined to the Paraná and Paraguay basins, or peripheral areas of the Gran Chaco ecoregion [80,108,113,114,115,116,117,118,119,120]. ...
The Gran Chaco is a wide ecologic-geographic region comprising northern Argentina, western Paraguay, southern Bolivia and the southwestern extreme of Brazil. This region exhibits extreme temperatures, annually regular frosts, and sedimentary soils; it has been dramatically threatened by agriculture expansion in recent decades. Therefore, increasing knowledge of plant diversity is critical for conservation purposes. We present a Legume checklist of the Gran Chaco ecoregion including conservation status of its endemic species. Leguminosae is the third most diverse plant family in the Neotropics. Assuming a rigorous spatial definition of the Gran Chaco, we recorded 98 genera, 362 species, and 404 specific and infraspecific taxa. Endemic/typical taxa were 17%, comparable to adjacent tropical plant formations, and they were found in higher percentages in Caesalpinioideae (24%) and Cercidoideae (33%) than Papilionoideae (11%) subfamily. We also analyzed the plant diversity comparing lineages and subregions. The Gran Chaco Legumes are predominantly widespread generalists, or they belong to either Chaco sensu stricto or Neotropical Seasonally Dry Tropical Forest (SDTF) lineages. Though the Humid Chaco registered the highest species richness, Dry Chaco and Sierra Chaco, the most threatrened subregions, exhibited the highest percentages of exclusive and proper Chaco-lineage species. These results suggest that diversification of Legumes has been most relevant in Dry Chaco and Sierra Chaco, probably by their more demanding and harsh environmental conditions limiting the dispersion of generalists or intrusive-invading species. This study is paramount to reach an improved delimitation of the Gran Chaco ecoregion in transitional areas with the SDTF and Cerrado formations. Conservation status is critical in genera of high economic interest, such as Arachis, Mimosa and Prosopis. At least one third of endemic taxa exhibit a critical status of conservation or are endangered, many of them being relevant to inbreeding program or exhibiting multiple economic uses.
... However, this approach was based on a single chloroplast clone of 499 bp in length, which might provide insufficient resolution to discriminate between S. seabrana and S. hamata. Moreover, the taxonomic status of S. seabrana (Maass and Mannetje, 2002) is dubious, since two recent studies have suggested that S. seabrana is a synonym for S. scabra (Vanni and Fernandez, 2011;Vanni, 2017). ...
Backgrounds and aims:
The genus Stylosanthes includes nitrogen-fixing and drought-tolerant species of considerable economic importance for perennial pasture, green manure and land recovery. Stylosanthes scabra is adapted to variable soil conditions, being cultivated to improve pastures and soils worldwide. Previous studies have proposed S. scabra as an allotetraploid species (2n = 40) with a putative diploid A genome progenitor S. hamata or S. seabrana (2n = 20) and the B genome progenitor S. viscosa (2n = 20). We aimed to provide conclusive evidence for the origin of S. scabra.
Methods:
We performed fluorescence in situ hybridization (FISH) and genomic in situ hybridization (GISH) experiments and Illumina paired-end sequencing of S. scabra, S. hamata and S. viscosa genomic DNA, to assemble and compare complete ribosomal DNA (rDNA) units and chloroplast genomes. Plastome- and genome-wide single nucleotide variation detection was also performed.
key results :
GISH and phylogenetic analyses of plastid DNA and rDNA sequences support that S. scabra is an allotetraploid formed from 0.63 to 0.52 million years ago (Mya), from progenitors with a similar genome structure to the maternal donor S. hamata and the paternal donor S. viscosa. FISH revealed a non-additive number of 35S rDNA sites in S. scabra compared with its progenitors, indicating the loss of one locus from A genome origin. In S. scabra, most 5S rDNA units were similar to S. viscosa, while one 5S rDNA site of reduced size most probably came from an A genome species as revealed by GISH and in silico analysis.
Conclusions:
Our approach combined whole-plastome and rDNA assembly with additional cytogenetic analysis to shed light successfully on the allotetraploid origin of S. scabra. We propose a Middle Pleistocene origin for S. scabra involving species with maternal A and paternal B genomes. Our data also suggest that variation found in rDNA units in S. scabra and its progenitors reveals differences that can be explained by homogenization, deletion and amplification processes that have occurred since its origin.
... However, this approach was based on a single chloroplast clone of 499 bp in length, which might provide insufficient resolution to discriminate between S. seabrana and S. hamata. Moreover, the taxonomic status of S. seabrana (Maass and Mannetje, 2002) is dubious, since two recent studies have suggested that S. seabrana is a synonym for S. scabra (Vanni and Fernandez, 2011;Vanni, 2017). ...
O gênero Stylosanthes (Fabaceae) possui grande importância agronômica por seu uso forrageiro e sua associação com bactérias nitrificantes, que o torna ideal para adubação verde e recuperação de solos degradados. O gênero tem o Brasil como principal centro de diversidade, sendo citogeneticamente conhecido por apresentar cariótipos estáveis com 2n = 20 e ocorrência de alotetraploides naturais, dentre esses S. scabra. Este trabalho teve como objetivo confirmar a origem desse alotetraploide a partir de S. hamata × S. viscosa. Para isso foi realizado o sequenciamento do DNA genômico total de S. scabra, S. viscosa e S. hamata utilizando a plataforma Illumina HiSeq 2000 com reads pareados. A caracterização da fração repetitiva dos genomas das três espécies foi feita individualmente e comparativamente no programa RepeatExplorer, no qual foi identificado a proporção das famílias de sequências repetitivas para cada espécie. Também foi feito a montagem do genoma plastidial (plastoma) das três espécies utilizando os programas NOVOPlasty v. 2.5.6 e Geneious v.9.1.8. A análise dos clusters obtidos no RepeatExplorer revelou vários grupos de DNA repetitivo presentes em S. scabra que eram específicos do genoma de S. hamata ou de S. viscosa, sendo esses principalmente pertencentes a famílias de DNA satélite. Uma análise dos contigs do DNA ribossomal (DNAr) mostrou alta similaridade de ambos DNAr 5S e 45S entre S. scabra e S. viscosa. A análise dos plastomas revelou uma alta similaridade (99,8%) entre o plastoma de S. scabra e S. hamata indicando que S. hamata seja seu possível parental materno. Análises de GISH corroboram essa relação de parentesco, mostrando que metade do complemento cromossômico de S. scabra é marcada com cada um do parentais S. hamata e S. viscosa. A análise comparativa dos sítios de DNAr 5S e 45S por hibridização in situ fluorescente (FISH) revela que S. scabra não possui o somatório dos sítios observados nos seus parentais, sugerindo algum grau de diploidização. A FISH das famílias de DNA satélite específicas e retroelementos de S. hamata e S. viscosa nos cromossomos do híbrido S. scabra poderá contribuir para o entendimento dos mecanismos de evolução e estabilização genômica desse alotetraploide.
Stylosanthes, with approximately 50 species, is a pantropical, economically important genus of Leguminosae, with several species having forage potential and soil improvement potential. Thirty-two species have been recorded in Brazil, with the Northeast region being one of the main centers of diversity of the genus in the country. Despite this, no taxonomic work focusing on Stylosanthes has been done for the region. This study presents a taxonomic treatment of Stylosanthes species occurring in the Northeast region of Brazil. Botanical specimens were collected in the field together with visits to the main herbaria in the Northeast (ALCB, HUEFS, HUNEB, HVASF) for specimen analysis. Online consultation of databases in virtual herbaria, analysis of regional flora accounts, assessment of types and discussions with legume experts were also undertaken. Twenty-three taxa were confirmed as occurring in NE Brazil, with six new occurrences recorded for the states of Ceará, Paraíba, Piauí, and Rio Grande do Norte. Species in the studied area are mainly differentiated by their fruit and bract morphology. Key words: campos rupestres, Cerrado, Fabaceae, flora, plant taxonomy, semiarid.
