Pod production, and dasometric variables, of the tree Senna spectabilis (Fabaceae) in a tropical dry forest / Producción de vainas, y variables dasométricas, del árbol Senna spectabilis (Fabaceae) en un bosque seco tropical

Introduction: Senna spectabilis is a multipurpose pantropical tree, used in agroforestry systems. Objective: To determine pod production (Pp) and their relationship with dasometric variables in S. spectabilis in the tropical dry forest. Methods: From August 2016 to February 2017, thirty trees in production stage were randomly selected. The random selection was formed of the more isolated trees from the total dispersion. The trees were monitored at the beginning and end of the study period, to determine dasometric measurements such as total height (Th), height to the first branch (Hb), crown height (Ch), Stem diameter (at 0.2 m height from the ground) (Db), crown diameter (Cd), and crown volume (Cv) measured. Pods were harvested by the researcher with cutting and height cutting tongs when their color began to change. Pearson correlations and univariate and multivariate regression analyses were performed between the dasometric variables and pod production. The potential number of trees/ha (NPa) was calculated by determining the occlusion percentage (Op) and the shadow area/tree (Ca); to estimate the production potential of fruits/ha, the production of fruits/tree was multiplied by (NPa). Results: Th was 6.16 ± 1.23 m, Hb 2.75 ± 0.52 m, Ch 3.41 ± 0.98 m, Db 20.43 ± 4.80 cm, Cd 7.46 ± 1.20 m and Cv 108.43 ± 61.38 m3/tree. There was a significant positive correlation between Hb, Cd, Db, with Pp of 0.592**, 0.592**, and 0.446* respectively. Pp was 32.73 ± 16.13 kg/tree and the dry matter production (MSP) was 17.84 ± 8.80 kg/tree. The result of the multivariate regression indicated that the second-order polynomial model presented best goodness of fit. Op was 73.4 7.92 %, the cup area was 49.3 m2/tree, Ca was 36.2 m2/tree, and NPa was 83 trees. Conclusions: The production of fresh pods/ tree in the S. spectabilis presents a potential in its availability as feed for ruminant or seed production. The potential production of pods in silvopastoral with S. spectabilis could be 2.72 t/ ha, and 1.64 t/ ha of dry pods, this shows the importance of trees and of pods production and nutritional contribution obtained for dry ecosystems.

Introducción: Senna spectabilis es un árbol pantropical multipropósito, utilizado en sistemas agroforestales. Objetivo: Determinar la producción de vainas (Pv) y la relación con las variables dasométricas en S. spectabilis en el bosque seco tropical. El número potencial de árboles/ha (NPa) fue calculado determinando el porcentaje de oclusión (Po) y el área de sombra/árbol (As); para calcular la producción potencial de frutos/ha, la producción de frutos/árbol fue multiplicada por (NPa). Métodos: Desde agosto del 2016 hasta febrero de 2017, treinta árboles en etapa de producción fueron seleccionados al azar, los más aislados del total de árboles dispersos fueron seleccionados, y fueron monitoreados al inicio y al final del período de estudio, para determinar las mediciones dasométricas como la altura total (At), altura a la primera rama (Apr), altura de la copa (Ac), diámetro del tallo (a 0.2 m altura desde el suelo) (Dt), diámetro de la copa (Dc) y volumen de copa (Vc). Las vainas se cosecharon cuando su color comenzó a cambiar. Se realizaron correlaciones de Pearson y análisis de regresión univariada y multivariada entre las variables dasométricas y la producción de vainas. El número potencial de árboles/ha (NPa) se calculó determinando el porcentaje de oclusión (Po) y el área de sombra/árbol (Asa); para estimar el potencial de producción de las vainas/ha, la producción de vainas/árbol se multiplicó por NPa. Resultados: La At fue de 6.16 ± 1.23 m, Apr 2.75 ± 0.52 m, Ac 3.41 ± 0.98 m, Db 20.43 ± 4.80 cm, Dc 7.46 ± 1.20 m y Vc 108.43 ± 61.38 m3/árbol. Existió una correlación positiva significativa entre Apr, Dc, Db, Pv de 0.592**, 0.592 ** y 0.446 * respectivamente. La Pv fue de 32.73 ± 16.13 kg y la producción de materia seca (PMS) fue de 17.84 ± 8.80 kg/árbol. El resultado de la regresión multivariada indicó que el modelo polinomial de segundo orden presentó la mejor bondad de ajuste. El Po de los árboles fue de 73.4 % ± 7.92 %, el área de copa fue de 49.3 m2/árbol, el Asa fue de 36.2 m2/árbol, el NPa fue de 83 árboles. Conclusiones: La producción de vainas frescas/árbol en el S. spectabilis presenta un potencial en la disponibilidad de alimento para los rumiantes o la producción de semillas. El potencial de producción de vainas en u arreglos silvopastoriles podría ser de 2.72 t/ha, y 1.64 t/ha de vainas secas, esto muestra la importancia del árbol de producción de vainas y la contribución nutricional para los ecosistemas secos.

Microscopic techniques used for the identification of medicinal plants: A case study of Senna.

BACKGROUND: The use of quality control tool for adulteration of Senna (Cassia aungustifolia) a pharmaceutically very important. They were used for multiple health disorders such as constipation, indigestion, epilepsy, asthma, piles, migraine, and heart problems. Two different species of same family or same genus used commercially in Indo-Pak using the same medicine name Senna. One named as Senna (C. aungustifolia) and its adulterant named as Sickle Senna (Cassia obtusifolia). METHODOLOGY: These two plants were analyzed using classical microscopic techniques light microscopy and the modern chemotaxonomic traits scanning electron microscopy, fluorescence studies and phytochemical studies. RESULTS: The C. aungustifolia L. had found to be a perennial herb with trilobed pollen, diacytic, paracytic, and anisocytic stomata having smooth walled epidermal cells, whereas the C. obtusifolia stands out as a perennial shrub with spheroidal and circular pollen and paracytic type of stomata having irregular shaped epidermal cells. The powdered drug of C. aungustifolia is dark grayish green, whereas the powdered drug of C. obtusifolia is light green in color. Investigation and other techniques used in this project provided the basis for the authentication of this species.

Heteranthery as a solution to the demand for pollen as food and for pollination - Legitimate flower visitors reject flowers without feeding anthers.

Heteranthery, the presence of feeding and pollinating anthers in the same flower, seems to mediate the evolutionary dilemma for plants to protect their gametes and yet provide food for pollinators. This study aims to elucidate the role of heteranthery in the buzz-pollinated Senna reniformis. The fecundity of pollen from long-, medium- and short-sized anthers was determined by hand cross-pollination experiments, and the quantity, size, ornamentation and viability of pollen of different anthers were compared. Rates of flower rejection by bees were measured in anther removal experiments to assess the preferences of flower visitors for feeding or pollinating anthers. Large bees, which were the effective pollinators of self-incompatible S. reniformis, avoided flowers without short feeding anthers, but not those without medium or long anthers. Illegitimate small and medium-sized bees were unresponsive to anther exclusion experiments. Long anthers deposited pollen on the back and short anthers on the venter of large bees. Pollen from long anthers had higher in vitro viability and higher fruit and seed set after cross-pollination than pollen from other sized anthers. Short anthers produce feeding pollen to effective pollinators and long anthers are related to pollination of S. reniformis. Bee behaviour and size was found to directly influence the role of anthers in the 'division of labour'. Only large bee pollinators that carry the pollinating pollen from long anthers in 'safe sites' associated short anthers with the presence of food. In the absence of these larger bee pollinators, the role of heteranthery in S. reniformis would be strongly compromised and its function would be lost.

Ricochet pollination in Senna (Fabaceae) - petals deflect pollen jets and promote division of labour among flower structures.

Naturalists Fritz and Hermann Müller hypothesised that heteranthery often leads to a division of labour into 'feeding' and 'pollinating' stamens; the latter often being as long as the pistil so as to promote successful pollination on the bees' back. In many buzz-pollinated species of Senna, however, the so-called pollinating stamens are short and not level with the stigma, raising the question of how pollen is shed on the bees' back. Here we explore a mechanism called 'ricochet pollination'. We test whether division of labour is achieved through the interaction between short lower stamens and strongly concave 'deflector petals'. We studied the arrangement and morphology of the floral organs involved in the ricochet pollination, functioning of the flowers through artificial sonication and observed the interactions between bees and flowers in the field. The middle stamens are adapted to eject pollen downwards, which can be readily collected on the bee mid legs. Most of the pollen is ejected towards the deflector petal(s). Pollen from this set of stamens is more likely to contribute to pollination. The pollen grains seem to ricochet multiple times against the deflector petals to eventually reach the bee's back. The pollen ricochet mechanism promotes a division of labour by involving additional floral organs, such as petals, reinforcing the Müllers' division-of-labour hypothesis. However, alternative, non-multiexclusive hypotheses could be explored in genus Senna and other angiosperm species.

Diversity and evolution of a trait mediating ant-plant interactions: insights from extrafloral nectaries in Senna (Leguminosae).

BACKGROUND AND AIMS: Plants display a wide range of traits that allow them to use animals for vital tasks. To attract and reward aggressive ants that protect developing leaves and flowers from consumers, many plants bear extrafloral nectaries (EFNs). EFNs are exceptionally diverse in morphology and locations on a plant. In this study the evolution of EFN diversity is explored by focusing on the legume genus Senna, in which EFNs underwent remarkable morphological diversification and occur in over 80 % of the approx. 350 species. METHODS: EFN diversity in location, morphology and plant ontogeny was characterized in wild and cultivated plants, using scanning electron microscopy and microtome sectioning. From these data EFN evolution was reconstructed in a phylogenetic framework comprising 83 Senna species. KEY RESULTS: Two distinct kinds of EFNs exist in two unrelated clades within Senna. 'Individualized' EFNs (iEFNs), located on the compound leaves and sometimes at the base of pedicels, display a conspicuous, gland-like nectary structure, are highly diverse in shape and characterize the species-rich EFN clade. Previously overlooked 'non-individualized' EFNs (non-iEFNs) embedded within stipules, bracts, and sepals are cryptic and may represent a new synapomorphy for clade II. Leaves bear EFNs consistently throughout plant ontogeny. In one species, however, early seedlings develop iEFNs between the first pair of leaflets, but later leaves produce them at the leaf base. This ontogenetic shift reflects our inferred diversification history of iEFN location: ancestral leaves bore EFNs between the first pair of leaflets, while leaves derived from them bore EFNs either between multiple pairs of leaflets or at the leaf base. CONCLUSIONS: EFNs are more diverse than previously thought. EFN-bearing plant parts provide different opportunities for EFN presentation (i.e. location) and individualization (i.e. morphology), with implications for EFN morphological evolution, EFN-ant protective mutualisms and the evolutionary role of EFNs in plant diversification.

Large-scale patterns of diversification in the widespread legume genus Senna and the evolutionary role of extrafloral nectaries.

Unraveling the diversification history of old, species-rich and widespread clades is difficult because of extinction, undersampling, and taxonomic uncertainty. In the context of these challenges, we investigated the timing and mode of lineage diversification in Senna (Leguminosae) to gain insights into the evolutionary role of extrafloral nectaries (EFNs). EFNs secrete nectar, attracting ants and forming ecologically important ant-plant mutualisms. In Senna, EFNs characterize one large clade (EFN clade), including 80% of its 350 species. Taxonomic accounts make Senna the largest caesalpinioid genus, but quantitative comparisons to other taxa require inferences about rates. Molecular dating analyses suggest that Senna originated in the early Eocene, and its major lineages appeared during early/mid Eocene to early Oligocene. EFNs evolved in the late Eocene, after the main radiation of ants. The EFN clade diversified faster, becoming significantly more species-rich than non-EFN clades. The shift in diversification rates associated with EFN evolution supports the hypothesis that EFNs represent a (relatively old) key innovation in Senna. EFNs may have promoted the colonization of new habitats appearing with the early uplift of the Andes. This would explain the distinctive geographic concentration of the EFN clade in South America.

Interspecific variation in vessel size, growth and drought tolerance of broad-leaved trees in semi-arid regions of Kenya.

In semi-arid regions, trees often wither during the dry season. Withering is sometimes manifest as die-back, whereby whithering results in shoot death, which progresses downward from the uppermost part of the crown. In this study, we measured the relationships between height growth and diameter at breast height, die-back frequency and severity, vessel size and specific hydraulic conductivity of four evergreen (Senna siamea (Lamk) H.S. Irwin & Barneby, Jacaranda mimosifolia D. Don, Azadirachta indica A.H.L. Juss and Acacia gerrardii Benth.) and one deciduous (Melia volkensii Gürke) plantation tree species in Kenya, which has a conspicuous dry season. Die-back occurred readily in some species, but not in others. Senna siamea showed the highest specific hydraulic conductivity and the highest growth rate among the five species and was quite susceptible to die-back. Among species, height growth and specific hydraulic conductivity were positively correlated with vessel size and negatively correlated with die-back frequency, suggesting a trade-off between growth rate and drought tolerance. This implies that an adaptation to rapid growth under humid conditions leads to low drought tolerance. However, the deciduous tree Melia volkensii showed high specific hydraulic conductivity and growth, with no symptoms of die-back, implying that a mechanism associated with the deciduous habit results in drought avoidance by reducing the requirement for water during the dry season.

Floral biology and reproductive system of enantiostylous Senna corymbosa (Caesalpiniaceae)

The genus Senna (K.Bahuin) Miller (Cassieae) is represented in Argentina by 35 species and several varieties distributed in temperate, tropical and subtropical regions, and presents a high degree of endemism. Some taxa are used for medicine, animal foraging and ornamental purposes. Floral morphology, phases, rewards, attractants, visitors, pollen, reproductive system, P/O ratio, OCI and ISI indexes of enantiostylous Senna corymbosa were analyzed for morphological androecial differentiation and possible related functional differences between stamens groups. The study was carried out over three consecutive flowering seasons in March of 1999, 2000 and 2001, in two populations near the city of Buenos Aires, Argentina. The species has a buzz-pollination syndrome. The pollinators are Bombus atratus that vibrate the stamens, leading to nothotribic or sternotribic deposition of pollen. P/O ratio and pollen production were high thanks to the precise pollinating mechanism that needs a large delivery of pollen to ensure effective pollen deposition. P/O ratio also indicates that the species is xenogamous, although geitonogamy and autogamy (only induced, not spontaneous)were also recorded. The species is self-compatible. There were differences in hand-pollination treatments between long and medium stamens in fruit set, as well as in vitro differential fertility between their pollen grains. Hence, there are morphological and functional androecium differences: the medium stamens play a nutritional role while the long ones play a reproductive one. These differences are reflected in the breeding system. Moreover, differential fertility and enantiostyly may diminish the effects of self-compatibility by partially reducing the contribution of geitonogamy to selfing

Se analizaron la morfología y fases florales, recompensa, atractivos, visitantes, polen, sistema reproductivo, índices OCI e ISI y relación polen/óvulo de la especie enantiostílica Senna corymbosa. El objetivo fue estudiar su biología floral y determinar la posible presencia de diferenciación morfológica en el androceo. Era importante saber si las diferencias morfológicas reflejaban diferencias funcionales entre los grupos de estambres. El estudio se llevó a cabo durante tres estaciones de floración consecutivas en marzo de 1999,2000 y 2001,sobre dos poblaciones presentes en zonas protegidas de la ciudad de Buenos Aires, Argentina. La especie presenta un síndrome de polinización vibrátil (buzz-pollination).Los polinizadores son individuos de Bombus atratus, capaces de vibrar los estambres y extraer polen, el que se deposita nototríbica y/o esternotríbicamente. La relación P/O fue extremadamente elevada, así como la producción de polen. Esto se relaciona con el comportamiento de los visitantes y con la alta precisión del mecanismo de polinización, que requiere una gran cantidad de polen para asegurar su efectiva deposición en el cuerpo del insecto. La relación P/O indica que la especie es xenógama, aunque se registraron también geitonogamia y autogamia inducidas. La especie, por lo tanto, es autocompatible. Se encontraron diferencias en los tratamientos de polinización manual entre estambres largos y medianos en cuanto al establecimiento de frutos, así como una fertilidad diferencial in vitro entre sus tipos polínicos. Existen, así, diferencias morfológicas y funcionales en el androceo: los estambres medianos tienen un papel nutricional mientras que los largos cumplen un papel reproductivo. Estas diferencias se reflejan en el sistema de cruzamiento. Más aún, la fertilidad diferencial y la enantiostilia disminuyen los efectos de la autocompatibilidad reduciendo parcialmente la contribución de la geitonogamia a la endogamia

Floral biology and reproductive system of enantiostylous Senna corymbosa (Caesalpiniaceae).

The genus Senna (K. Bahuin) Miller (Cassieae) is represented in Argentina by 35 species and several varieties distributed in temperate, tropical and subtropical regions, and presents a high degree of endemism. Some taxa are used for medicine, animal foraging and ornamental purposes. Floral morphology, phases, rewards, attractants, visitors, pollen, reproductive system, P/O ratio. OCI and ISI indexes of enantiostylous Senna corymbosa were analyzed for morphological androecial differentiation and possible related functional differences between stamens groups. The study was carried out over three consecutive flowering seasons in March of 1999, 2000 and 2001, in two populations near the city of Buenos Aires, Argentina. The species has a buzz-pollination syndrome. The pollinators are Bombus atratus that vibrate the stamens, leading to nothotribic or stemotribic deposition of pollen. P/O ratio and pollen production were high thanks to the precise pollinating mechanism that needs a large delivery of pollen to ensure effective pollen deposition. P/O ratio also indicates that the species is xenogamous, although geitonogamy and autogamy (only induced, not spontaneous) were also recorded. The species is self-compatible. There were differences in hand-pollination treatments between long and medium stamens in fruit set, as well as in vitro differential fertility between their pollen grains. Hence, there are morphological and functional androecium differences: the medium stamens play a nutritional role while the long ones play a reproductive one. These differences are reflected in the breeding system. Moreover, differential fertility and enantiostyly may diminish the effects of self-compatibility by partially reducing the contribution of geitonogamy to selfing.