M.M. Alemán’s research while affiliated with National University of Salta and other places

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Publications (15)


Flowers of the studied species. aCologania broussonetii; bCrotalaria chaco-serranensis; cC. pumila; dC. stipularia; eDesmodium incanum; fD. pachyrrhizum; gD. subsericeum;hD. uncinatum; iGalactia latisiliqua; jIndigofera parodiana; kI. suffruticosa, lMacroptilium erythroloma; mM. gibbosifolium; nPhaseolus vulgaris var. aborigineus; oRhynchosia edulis; pR. minima; qZornia contorta
Schematic representation of the main pollination mechanisms in Papilionoideae and quantitative traits that were measured in this study. a valvular; b explosive; c Brush; d pump; e quantitative measured traits (fs flower size, gl gynoecium length, k keel, kc keel claw length, kl keel length, kw keel width, ls longest stamen length, sl style length, ss shortest stamen length, scl staminal column length, scw staminal column width, v vexillum, vl vexillum length, vw vexillum width, vc vexillum claw length, w wings, wc wing claw length, wl wing length, ww wing width; arrows indicate the movement of the floral pieces during the activation of the mechanisms; scale bars in a = 1.6 mm, b = 1.2 mm, c = 0.8 mm, d = 1.2 mm, e = 1 mm)
Scatter plots a PC1 and PC2; b PC1 and PC3; c PC2 and PC3. (C.inc Crotalaria chaco-serranensis; C.pum C. pumila; C.sti C. stipularia; Colog Cologania broussonetii; D.inc Desmodium incanum; D.pac D. pachyrrhizum; D.sub D. subsericeum; D.unc D. uncinatum; Galac Galactia latisiliqua; I.par Indigofera parodiana; I.suf I. suffruticosa; M.ery Macroptilium erythroloma; M.gib. M. gibbosifolium; Phase Phaseolus vulgarisvar.aborigineus; R.edu Rhynchosia edulis; R.min R. minima; Zorn Zornia contorta)
Alemán M.M., Hoc P., Etcheverry A.V., Ortega-Baes P., Sühring S. y Lopez-Spahr D. 2022. Morphological traits in keel flowers of Papilionoideae (Fabaceae) and their relationships with the pollination mechanisms. Plant Systematics and Evolution. [in press] doi: 10.1007/s00606-022-01826-y.
  • Article
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October 2022

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205 Reads

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1 Citation

Plant Systematics and Evolution

María Mercedes Alemán

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Patricia Hoc

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Several floral traits in Papilionoideae (Fabaceae) have been correlated to certain pollination mechanisms. To test this hypothesis, we analyzed 20 qualitative and 21 quantitative floral traits of 17 species of Papilionoideae with keel flowers from various tribes and genera native to Salta, Argentina, which are representative of the four general pollination mechanisms described for the subfamily. We performed field and laboratory studies, including description of morphology, micro-morphology, position, size, and synorganization of all floral organs. Correlation between floral traits and a particular pollination mechanism was evaluated through a principal component analysis (PCA). According to our results, species with the same pollination mechanism share similar qualitative and quantitative traits, the latter being the most relevant, according to our statistical analysis.

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Fig. 1. Vista esquemática de una flor típica de Papilionoideae mostrando la ubicación de las piezas florales. La ilustración corresponde a Crotralaria chaco-serranensis. Colores= verde: cáliz, amarillo: vexilo, naranja: ala, rosa: quilla, rojo: androceo, azul: gineceo.
Fig. 5. Vista al MEB de los estigmas. A: Cologania broussonetii. B: Crotalaria chaco-serranensis. C: Detalle de B (estigma sin membrana, se observan las papilas estigmáticas). D: C. pumila. E: Desmodium subsericeum. F: Galactia latisiliqua. G: Indigofera suffruticosa. H: Macroptilium gibbosifolium. I: M. erythroloma. J: Phaseolus vulgaris var. aborigineus. K: Rynchosia edulis. L: Zornia contorta. M: R. edulis (en flor sin visitar). N: detalle de M (se observa la membrana desprendida espontáneamente). O: D. incanum (en flor sin visitar). P: detalle de O (se observa una fisura de la membrana). Los recuadros indican la zona ampliada.
Diversidad morfológica del androceo y gineceo de 17 especies de Papilionoideae (Fabaceae) nativas del Valle de Lerma (Salta-Argentina) y su relación con algunos aspectos eproductivos

September 2020

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104 Reads

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1 Citation

Boletin de la Sociedad Argentina de Botanica

Introducción y objetivos: Las especies de Papilionoideae presentan características florales o estrategias relacionadas con los mecanismos de polinización y que favorecen la polinización cruzada como la presencia de barreras físicas, la separación espacial o temporal en la maduración de los ciclos fértiles, entre otras. El objetivo de este trabajo fue brindar una descripción detallada de la morfología del androceo y gineceo de 17 especies de Papilionoideae nativas del Valle de Lerma (Salta-Argentina). M&M: Se estudiaron a campo y en laboratorio características cuantitativas y cualitativas del androceo y el gineceo en flores de 17 especies de Papilionoideae, pertenecientes a nueve géneros y cinco tribus y se analizaron desde un punto de vista funcional aquellas características que estarían relacionadas con el sistema reproductivo y los mecanismos de polinización. Resultados: Los resultados indican que la mayoría de las especies son homógamas, presentan androceo monadelfo o pseudomonadelfo con fenestras basales, anteras monomórficas, formas variables del estilo y del estigma, tricomas estilares y periestigmáticos y membrana estigmática. Se registraron características descritas por primera vez para algunas de estas especies tales como la ruptura espontánea de la membrana estigmática durante la antesis y la reducción de la distancia anteras-estigma luego de la maduración de los ciclos sexuales. Conclusiones: Todas las especies estudiadas presentan características florales asociadas a la reducción de la autopolinización. Se obtuvo información detallada sobre la morfología y otras características del androceo y gineceo que podría ser utilizada para describir mejor sus mecanismos de polinización y para comprender aspectos de su biología reproductiva.


Fusión, esculturas y ornamentaciones de las piezas de la corola de 17 especies de Papilionoideae

December 2017

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783 Reads

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1 Citation

Boletin de la Sociedad Argentina de Botanica

The species of subfamily Papilionoideae present flowers with a complex architecture, with elaborated petals in which it has been documented the presence of auricles and callosities, appendixes or spurs that joint the pieces of the corolla and also microsculpturings on the wings. In this study, we analyze the morphology of the petals of 17 species of Papilionoideae (Leguminosae) native of Salta province, Argentina. The lower margins of the keel were fused in all of the species, through different types of fusion of epidermal cells. Only a few species presented fusion in the upper margin of the keel. All the species showed sculpturings in the petals, which would joint some of the pieces of the corolla. They also exhibit ornaments or microsculpturings in the basal region of the adaxial surface of the wings that may act as a grip for pollinators when they land on the flower. The petals of the Papilionoideae flowers presented characteristics that could restrict the type of pollinator able to activate the different mechanisms of pollination that allow them to access the rewards enclosed by the keel.


Fig. 1 Studied species. a Desmodium subsericeum, b D. uncinatum and c D. incanum  
Fig. 2 a Detail of a wing petal, showing folds in the shallow folder. b Detail of the folds. c Wing–keel complex (1: wing depression, 2: keel protuberance). d Cuticle connection between the wing and keel.  
Fig. 3 Floral traits of Desmodium subsericeum. a Different floral stages (1: bud, 2: flower in anthesis without the explosive mechanism activated, 3: flower in anthesis with the mechanism activated). b Staminal column, showing the position of the stamens. c Pollen  
The explosive pollination mechanism in Papilionoideae (Leguminosae): An analysis with three Desmodium species

January 2014

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2,669 Reads

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33 Citations

Plant Systematics and Evolution

In the papilionaceous flowers with explosive pollination mechanism, the androecium and the gynoecium are subjected to great pressure in the keel. It has been suggested that at the first visit nearly all pollen grains are released and that the floral parts do not return to their original positions; this would be a cue for pollinators to avoid activated flowers. The dependence on the pollinators suggests that explosive flowers will not produce fruits if the mechanism is not activated. In this paper, we have tested these ideas studying the reproductive biology of three Desmodium species, D. incanum, D. subsericium and D. uncinatum. Specifically, we studied the floral traits, reproductive system, pollination mechanism and floral visitors, for the three species. The results indicated that the keel and the wings play a key role in the activation of the explosive mechanism. The three species have a secondary pollen presentation, diurnal flowers, and a short floral cycle. According to the results, all the species studied were self-compatible, with two species producing fruits by autonomous self-pollination. Desmodium flowers were visited and activated by native and exotic bee species. During the first visit nearly all the pollen was released; after that, new visits were low or null. Our results indicated that the activation of the mechanism is not essential for the production of fruits, that explosive flowers are visited only once and that the changes in the positions of the floral parts are the cues that keep pollinators away from the flowers with activated mechanism.



Generalised pollination system of Erythrina dominguezii (Fabaceae : Papilionoideae) involving hummingbirds, passerines and bees

September 2012

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1,534 Reads

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20 Citations

Plants in the genus Erythrina are pollinated by birds, such that passerine pollination is the plesiomorphic state, whereas hummingbird pollination is the derived character. Phylogenetic studies suggest that Erythrina dominguezii belongs to a basal clade characterised as pollinated by both passerines and hummingbirds. Here, we characterise the pollination system of E. dominguezii. Floral morphology, nectar traits, breeding system, visitation rates and pollen deposition by its floral visitors were studied. Floral morphology of E. dominguezii showed traits associated with both passerine and hummingbird pollination. Nectar sugar concentration showed an intermediate value but closer to the hummingbird type; however, it was rich in hexose, which is typical of the passerine type. Approximately 5% of the flowers set fruits under free pollination. Almost 80% of recorded flowers were visited by birds, with the rest visited by hymenopterans (bumblebees and honeybees). Among avian pollinators, five species of hummingbirds and three passerine species were identified as pollinators. The hummingbird Chlorostilbon lucidus was the most efficient visitor in terms of pollen deposition and was second in frequency of visits. The passerine Icterus cayanensis was second in efficiency at depositing pollen and was the most frequent pollinator. Our results show that E. dominguezii has a generalised pollination system. In addition, we report a new case of closed flowers and secondary nectar presentation. This is the first study that compares effectiveness among different pollinators in Erythrina.


Biodiversity of medicinal plants from salta province, Northwestern Argentina

August 2012

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33 Reads

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1 Citation

Acta Horticulturae

Argentina is one of the most diverse countries in the world due to its species richness and endemism. A survey was carried out to categorize medicinal Angiosperms weeds found in Salta Province. We found that Dicotyledones constitutes 92.1%, with 65 families and 232 genera. Asteraceae turned out to be the family with the largest number of used taxa (24.55% of the total medicinal plants), followed by Fabaceae (9.02%), Solanaceae (7.67%), Verbenaceae (6.65%) and Lamiaceae (5.63%). Monocotyledones are represented by 10 families and 26 genera. The most important families were Poaceae (4.16% of the total medicinal plants), Typhaceae (1.04%), Liliaceae (0.52%) and Cannaceae (0.52%). The recorded species were associated with 33 health problems. Considering the most common medicinal uses we recorded that 17% of the species had an antibiotic activity, 9.32% of the species had antifungal activity, and 7.6% had anti-inflammatory activity. The most frequently used plant part were leaves (25.6%), stems (15.9%) and roots (10.6%). These taxa are still used for the prevention and relief of medical disorders. The number of species covered in this review demonstrates that Salta Province represents an important source of medicinal plants.


Generalised pollination system of Erythrina dominguezii (Fabaceae : Papilionoideae) involving hummingbirds, passerines and bees

January 2012

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530 Reads

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6 Citations

Plants in the genus Erythrina are pollinated by birds, such that passerine pollination is the plesiomorphic state, whereas hummingbird pollination is the derived character. Phylogenetic studies suggest that Erythrina dominguezii belongs to a basal clade characterised as pollinated by both passerines and hummingbirds. Here, we characterise the pollination system of E. dominguezii. Floral morphology, nectar traits, breeding system, visitation rates and pollen deposition by its oral visitors were studied. Floral morphology of E. dominguezii showed traits associated with both passerine and hummingbird pollination. Nectar sugar concentration showed an intermediate value but closer to the hummingbird type; however, it was rich in hexose, which is typical of the passerine type. Approximately 5% of the owers set fruits under free pollination. Almost 80% of recorded owers were visited by birds, with the rest visited by hymenopterans (bumblebees and honeybees). Among avian pollinators, ve species of hummingbirds and three passerine species were identied as pollinators. The hummingbird Chlorostilbon lucidus was the most efcient visitor in terms of pollen deposition and was second in frequency of visits. The passerine Icterus cayanensis was second in efciency at depositing pollen and was the most frequent pollinator. Our results show that E. dominguezii has a generalised pollination system. In addition, we report a new case of closed owers and secondary nectar presentation. This is the rst study that compares effectiveness among different pollinators in Erythrina.


Pollen:ovule ratio and its relationship with other floral traits in Papilionoideae (Leguminosae): An evaluation with Argentine species

January 2012

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238 Reads

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40 Citations

Plant Biology

The pollen:ovule ratio (P/O) has traditionally been used as a rough estimator of plant breeding systems. It has been shown that plant breeding systems are associated with particular floral traits. In this study, we determined the P/O in 21 Leguminosae species from Argentina and explored relationships between P/O and taxonomic position, flower size, floral rewards, pollen presentation and pollination mechanisms. According to the results, 15 out of the 21 species classified were obligate xenogamous, although some of them have been recorded as facultative xenogamous in previous studies. There was a significant effect of taxonomic position (genus), reward type and pollination mechanism on P/O. Species offering only nectar as a floral reward (which were species with a brush mechanism) had a significantly lower P/O than species offering pollen or pollen and nectar. Species with the brush pollination mechanism had the lowest P/O, while species with valvular and pump mechanism had the highest P/O. However, pollen presentation (primary and secondary) and flower size did not have a significant effect on P/O. Our results demonstrate that P/O variability is determined by taxonomic position and pollination mechanism in this plant group.


Growth habit and root anatomy in Macroptilium erythroloma and Macroptilium bracteatum (Fabaceae)

January 2012

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45 Reads

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4 Citations

We studied the growth habit and root anatomy on two species of the Macroptilium genera, Fabaceae. These species, potential forages, are native to the Northwest Provinces in Argentina. The aims of this paper were to determine the survival capacity of the study species relating its growth habit and persistence with morphological and anatomical features. Root samples were collected and cut using either a microtome or by hand, at varying distances from the root tip. Cuts were examined under a light microscope, and with scanning electron microscopy (SEM) using staining techniques. Macroptilium bracteatum and M. erythroloma are perennial herbs. Aerial parts die during winter, and sprouting occurs in spring from the innovation area, throughout sub-superficial buds. In addition, M. bracteatum and M. erythroloma develop root succulence as a strategy of persistence. Besides, M. bracteatum presents endogenous root buds arising from one-year-old roots with secondary growth. These results indicate that both Macroptilium bracteatum and M. erythroloma have good potential as forage species.


Citations (11)


... keel (with or without torsion, i.e. the keel can be lengthened and tubular in appearance forming one or more superimposed revolutions constituting a spiral, Tucker 1989; Yeo 1993), (6) fusion of the keel petals (along lower or upper margins or complete), (7) type of connection between wings and keel (without connection, with morphological connection, with morphological and epidermal connection; following Alemán et al. (2017), occurs when there is a protrusion of the abaxial surface of the keel and a depression in the adaxial surface of the wings; a morphological and cuticular connection occurs when the petals have morphological connection and the epidermis in contact are fused at cuticular level), (8) presence of sculpturing in the vexillum, (9) presence of sculpturing in the wings (auricles, appendages, or spurs and depressions), (10) presence of sculpturing in the keel, (11) presence of wing ornamentations (microsculptures in the epidermis as in Alemán et al. 2017), (12) concrescence of the staminal filaments (monadelphous, pseudo-monadelphous and diadelphous), (13) shape of the anthers of the outer stamens (oblong, orbicular, suborbicular, globose), (14) shape of the anthers of the inner stamens, (15) type of insertion of the filament in the anthers of the outer stamens (basifixed, dorsifixed, apicifixed), (16) type of insertion of the filament in the anthers of inner stamens, (17) shape of the style (linear, linear curved towards the apex, linear curved towards the base, linear angulated, linear curved (20) presence of stigmatic papillae covered by cuticle (following Alemán et al. 2020). The pollination mechanisms considered were brush (BM), pump (PM), valvular (VM) and explosive (EM). ...

Reference:

Alemán M.M., Hoc P., Etcheverry A.V., Ortega-Baes P., Sühring S. y Lopez-Spahr D. 2022. Morphological traits in keel flowers of Papilionoideae (Fabaceae) and their relationships with the pollination mechanisms. Plant Systematics and Evolution. [in press] doi: 10.1007/s00606-022-01826-y.
Diversidad morfológica del androceo y gineceo de 17 especies de Papilionoideae (Fabaceae) nativas del Valle de Lerma (Salta-Argentina) y su relación con algunos aspectos eproductivos

Boletin de la Sociedad Argentina de Botanica

... Resprouting in M. lathyroides plants starts from vegetative buds. In some Macroptilium species, resprouting occurs throughout sub-surface vegetative buds and roots accumulate reserves (carbohydrates) that provide the necessary energy to promote regrowth (Montiel et al. 2012), which probably occurs in M. lathyroides. The xylopodium (woody subterranean organ that is sometimes slightly fleshy, derived from the roots) is present in many perennial herbaceous legumes, which enables plants to persist during adverse seasons when aerial parts disappear (Burkart 1952). ...

Hábito de crecimiento y anatomía de la raíz en Macroptilium erythroloma y Macroptilium bracteatum (Fabaceae)
  • Citing Article
  • December 2012

Phyton-International Journal of Experimental Botany

... iv. Macroptilium bracteatum CPI 55770: a perennial species, to represent the genus Macroptilium; found to be the most outstanding perennial species for use in ley pastures in central Queenslandtwo very similar accessions have been released as cultivars cv Cadarga (CPI 55769) and cv Juanita (CPI 68892) (Pengelly, et al., 1998); and presents endogenous root buds from one-year-old roots with secondary growth (Martin Montiel et al., 2012). v. Arachis triseminalis CPI 91423: a survivor in north Queensland's rangelands (Amar and Gardiner, 1999). ...

Growth habit and root anatomy in Macroptilium erythroloma and Macroptilium bracteatum (Fabaceae)
  • Citing Article
  • January 2012

... 9) is not commonly found in floral volatiles, rarely above 1 % excepted in inflorescence of Vigna caracalla (L.) Verdc. [24] and Rosa x damascena 'Celsiana', [25] its relative content in the headspace E. submacrophylla flowers should be underlined, offering a singular olfactory facet to the "white-floral" note. Likewise, methyl nicotinate (no. ...

Characterization of Vigna caracalla fragrance
  • Citing Article
  • February 2010

Acta Horticulturae

... results). All the studied species are of great interest to landscape gardening (Etcheverry et al., 2010) and may also be used for forage given their high nutritional value (Fernández et al., 1988). ...

Fabaceae from Northwestern Argentina and their potential use as ornamentals
  • Citing Article
  • February 2010

Acta Horticulturae

... En particular, esta familia es la más abundante en número de especies medicinales en el Altiplano. En la provincia de Salta, Argentina, se encontró que las familias con mayor número de taxones usados, sobre el total de plantas medicinales, fueron Asteraceae (35.9%), seguida por Fabaceae (13.3%) (Etcheverry et al. 2012). Asimismo, Barbarán (2018) identificó 42 especies de plantas medicinales en Salta y Jujuy, entre 3500 y 5000 m s. n. m.; entre ellas, Asteraceae es la familia con más especies (30.9%). ...

Biodiversity of medicinal plants from salta province, Northwestern Argentina
  • Citing Article
  • August 2012

Acta Horticulturae

... Plants from many different families have evolved convergent mechanisms to ballistically release their pollen, such as Fabaceae (Raju and Rao 2006;Alemán et al. 2014), Lamiaceae (Harley 1971;Aluri 1990), Marantaceae (Davis 1987), Loranthaceae (Feehan 1985), and Hydrocharitaceae (Zhang 2020). In these plants, all or most of a flower's pollen is fired at the pollinator when it visits a flower for the first time. ...

The explosive pollination mechanism in Papilionoideae (Leguminosae): An analysis with three Desmodium species

Plant Systematics and Evolution

... When located in the vegetative organs, they generally provide recompenses for animals, mainly ants, which confer them protection against herbivorous predation and pathogens attacks (Heil, 2011;Beaumont et al., 2016). If present in the flower, the produced nectar often serves as a reward for pollinators (Heil, 2011;Etcheverry et al., 2012). ...

Generalised pollination system of Erythrina dominguezii (Fabaceae : Papilionoideae) involving hummingbirds, passerines and bees

... These floral traits affect the type of pollinators and their activity (Arroyo 1981; Tucker 1987, Westerkamp 1997 Westerkamp and Weber 1999). The flowers of this plant group are more commonly visited by bees (Arroyo 1981; Westerkamp 1997; Westerkamp and Weber 1999), although birds (Arroyo 1981; Etcheverry and TruccoAlemán 2005; Agostini et al. 2006; Etcheverry et al. 2012b), lizards (Sazima et al. 2005), and mammals (Baker 1970; Tandon et al. 2003) have also been reported as pollinators. In papilionaceous flowers, pollen deposition and removal can occur through four types of specific pollination mechanisms: valvular, pump, explosive and brush (Arroyo 1981; Yeo 1993; Westerkamp 1997; Westerkamp and Weber 1999; López et al. 1999; Galloni et al. 2007; Etcheverry et al. 2012a ). ...

Generalised pollination system of Erythrina dominguezii (Fabaceae : Papilionoideae) involving hummingbirds, passerines and bees

... We now confront an empirical pattern: species with more efficient pollen transfer from donor anther to receptive stigma tend to produce fewer pollen grains per ovule (Cruden, 2000;Gong & Huang, 2014;Pellmyr et al., 2020). Surrogate measures of pollen transfer efficiency, such as morphological features that enable precise placement of pollen or more effective pollen removal by stigmas (Cruden, 2000;Chouteau et al., 2006;Galloni et al., 2007;Etcheverry et al., 2012) or ecological circumstances thought to provide more reliable pollination (Cunha et al., 2022), are also associated with lower pollen-ovule ratios. If, as our model suggests, such associations are not causal relationships, how do they arise? ...

Pollen:ovule ratio and its relationship with other floral traits in Papilionoideae (Leguminosae): An evaluation with Argentine species
  • Citing Article
  • January 2012

Plant Biology