Resumen
Anadenanthera colubrina var. cebil is the paradigmatic tree species of the Seasonally Dry Tropical Forest (SDTF) of central South America. The southernmost part of its range is divided into two disjunct nuclei: Misiones (Mi) and the Sub-Andean Piedmont (SAP), separated by some 800 km and with contrasting environmental conditions. This disjunct distribution provides a suitable experimental scenario to evaluate possible adaptive changes. We sampled 262 individuals of the species from 26 natural populations from both nuclei and two rare ones from the Humid Chaco. Fresh leaves and ripe fruits were collected from each tree, on which 13 foliar and seven reproductive morphological traits were measured. Differences between regions and variability between and within populations were analysed using a linear mixed factorial model. To determine the possible relationship between phenotypic variability of morphological traits and ecological environmental variables, a redundancy analysis (RDA) was performed considering climate data from the CHELSA database, the enhanced vegetation index, and main soil depth. The three regions differed in half of the traits studied, with the greatest contrast between SAP and Mi, while the Humid Chaco was most similar to the latter. Variability between and within populations was found in almost all traits studied, indicating possible adaptive effects in the past and the potential occurrence of adaptations in the future. The ‘tree’ factor showed greater variability than the ‘population’ factor for almost all traits and even explained more of the variance than the residuals of the model. The RDA showed a clear separation between SAP and Mi populations and identified isothermality, mean temperature of the driest quarter, seasonality of precipitation, and precipitation of the warmest quarter as associated with the variability of the traits considered. These results are evidence of adaptation and also of a common gene pool due to a shared evolutionary history of both SDTF nuclei. Genetic studies using molecular markers and common garden trials should be carried out to gain more reliable conclusions.
Métodos
Plant material In total, 262 individuals belonging to 28 populations of A. colubrina var. cebil were sampled in the Misiones and Sub-Andean Piedmont SDTF nuclei and in the Humid Chaco. Several voucher specimens were added to the herbarium of the Museo de Ciencias Naturales de Salta (Horco Molle population: García and Barrandeguy 38—MCNS11943, García and Barrandeguy 37—MCNS11946; Calilegua population: García and Barrandeguy s.n.—MCNS11941, García and Barrandeguy s.n.—MCNS11942; Puerto Dalmacia population: García and Barrandeguy s.n.—MCNS 11947; Candelaria population: Barrandeguy and García s.n.—MCNS 11951, Barrandeguy and García s.n.—MCNS 11948; Santa Ana population: Barrandeguy and García s.n.—MCNS 11949, Barrandeguy and García s.n.—MCNS 11950). Additionally, three to five leaves from each sampled tree are preserved in the herbarium of the Department of Genetics—Facultad de Ciencias Exactas, Químicas y Naturales, Universidad Nacional de Misiones (Argentina), available for future studies. Note that the markedly unequal number of populations sampled in each region reflects the unequal distribution and relative abundance of the species in these three regions. The geographical location of the populations is detailed in Table 1. From each tree, at least five fresh and healthy leaves were collected from branches in the centre of the canopy, which were dried and stored. Ripe fruits were also collected from most of the trees. Analysed traits Five leaves and between two and 88 fruits per individual were measured. Seed traits were measured on all seeds of the analysed fruits. All measured leaves, fruits, and seeds were photographed with a Nikon D5500 digital camera and processed with the ImageJ software. The study is based on 20 morphological traits: 13 leaf traits and seven reproductive traits (in fruits and seeds). The leaf traits analysed were: petiole length (PeL), basal pinna length (BPL), median pinna length (MPL), terminal pinna length (TPL), basal pinna width (BPW), median pinna width (MPW), terminal pinna width (TPW), distance between pairs of median pinnae (MPD), leaf blade length (LBL), leaf blade width (LBW), number of pairs of pinnae (PPN), number of pinnulas in terminal pinnae (PiN), and leaf area (LeA). The analysed reproductive traits were: length (FL), width (FW), and area (FA) of the fruits, number of locules per fruit (LN), and length (SL), width (SW), and area (SA) of the seeds. The traits BPL, BPW, MPL, MPW, TPL, and TPW indicate the average between the pinnae of each pair. MPD was measured between the insertions of pairs of median pinnae with an intermediate pair on leaves with an odd number of pinnae, and without an intermediate pair on leaves with an even number of pinnae. The values of FA and SA were measured in 15 of the 28 populations, according to the availability of the material.