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Revista Chapingo serie ciencias forestales y del ambiente

versión On-line ISSN 2007-4018versión impresa ISSN 2007-3828

Rev. Chapingo ser. cienc. for. ambient vol.26 no.3 Chapingo sep./dic. 2020  Epub 25-Jun-2021

https://doi.org/10.5154/r.rchscfa.2019.12.092 

Scientific note

Litterfall from tropical dry forest trees scattered in pastures

Ma. de la Luz Avendaño-Yáñez1 

Salome Quiroz-Martínez2 

Sergio Pérez-Elizalde3 

Silvia López-Ortiz1  * 

1Colegio de Postgraduados, Campus Veracruz. km 88.5 Carretera Federal Xalapa-Veracruz, Predio Tepetates. C. P. 91690. Manlio Fabio Altamirano, Veracruz, México.

2Conselva, Costas y Comunidades A. C. Av. Olas Altas núm. 66, col. Centro. C. P. 82000. Mazatlán, Sinaloa, México.

3Colegio de Postgraduados, Campus Montecillo. km 36.5 Carretera Federal México-Texcoco, Montecillo. C. P. 56230. Texcoco, Estado de México, México.


Abstract

Introduction:

Scattered trees from tropical dry forests (TDF) deposit significant amounts of leaf litter into pasture soils.

Objective:

To estimate the litterfall production during the dry season by scattered Lysiloma acapulcense (Kunth) Benth. and Vachellia pennatula (Schltdl. & Cham.) Seigler & Ebinger trees in pastures.

Materials and methods:

Ten scattered trees of each species were selected from a 10-ha pasture, and litter was collected during the dry season (November to May) and sorted into fractions. Leaf litter biomass, nitrogen (N) and lignin (L) content, and the lignin-nitrogen (L/N) ratio were assessed. Leaf litterfall was modelled over an annual cycle using a double logistic model. Means from all variables were compared between species using a t-test.

Results and discussion:

Litter, nitrogen and lignin content were similar between species. However, the L/N ratio was significantly higher (P < 0.0001) in L. acapulcense (32.1) than in V. pennatula (21.3). The leaf litter model showed that the defoliation phase of L. acapulcense lasted 30 days longer than that of V. pennatula.

Conclusion:

The quantity and quality of leaf litter that scattered trees deposit in a pasture encourages the adoption of agroforestry systems, including tree species native to TDF.

Keywords: Vachellia pennatula; Lysiloma acapulcense; lignin; nitrogen; legume species

Resumen

Introducción:

Los árboles dispersos del bosque tropical seco (BTS) depositan una cantidad significativa de hojarasca en suelos de los pastizales.

Objetivo:

Estimar la producción de hojarasca de árboles dispersos de Lysiloma acapulcense (Kunth) Benth. y Vachellia pennatula (Schltdl. & Cham.) Seigler & Ebinger en pastizales, durante la estación seca.

Materiales y métodos:

Se recolectó la hojarasca de 10 árboles dispersos de cada especie en un pastizal de 10 ha, durante el periodo de senescencia de las hojas (noviembre a mayo). La cantidad de hojarasca se evaluó y se separó en componentes para determinar el contenido de nitrógeno (N) y lignina (L), y la proporción L/N. La caída de hojas se analizó con un modelo logístico doble. Las medias de las variables se compararon entre especies mediante una prueba t de Student.

Resultados y discusión:

La cantidad de hojarasca y los contenidos de N y L no difirieron entre especies (P > 0.05), mientras que la proporción L/N fue mayor en L. acapulcense (32.1) que en V. pennatula (21.3). La fase de defoliación de L. acapulcense es más larga que la de V. pennatula.

Conclusión:

Es importante conocer la cantidad y la calidad de la hojarasca que los árboles dispersos aportan en los pastizales para ayudar a la adopción de sistemas agroforestales, que incluyan especies nativas del BTS.

Palabras clave: Vachellia pennatula, Lysiloma acapulcense, lignina; nitrógeno, leguminosas

Introduction

Scattered trees are predominant components of many anthropogenic landscapes, particularly in livestock grazing systems (Rivest, Paquette, Moreno, & Messier, 2013). In pastures, scattered trees increase soil organic carbon and nutrients (Aryal, Gómez-González, Hernández-Nuriasmú, & Morales-Ruiz, 2018; Avendaño-Yáñez, López-Ortiz, Perroni, & Pérez-Elizalde, 2018) through litter accumulation. For example, scattered Lysiloma acapulcense (Kunth) Benth. and Vachellia pennatula (Schltdl. & Cham.) Seigler & Ebinger trees increase the nitrogen and phosphorus content in tropical pasture soils (Avendaño-Yáñez et al., 2018). Both species are native to tropical dry forest (TDF), and are used as multipurpose trees in agroecosystems: L. acapulcense is used as shade for livestock, as a source of timber for construction, fencing and firewood; V. pennatula is considered as a weed, even though its forage fruits and firewood are widely used (Suárez et al., 2012; Williams-Linera, Alvarez-Aquino, Hernández-Ascención, & Toledo, 2011). Yet, the ecological traits and ecosystem services of these two tree species, such as litterfall and its role in soil nutrient inputs, are not fully understood.

Leaf quantity and quality are linked to nutrient cycling in terrestrial ecosystems (Dutta & Agrawal, 2001), because leaves are the primary source of organic matter for the soil (Sayer & Tanner 2010; Vitousek & Sanford, 1986). Leaf litter decomposition depends on its chemical composition (Loranger, Ponge, Imbert, & Lavelle, 2002), mainly lignin content, because it controls the decomposition rate of plant residues (Austin & Ballaré, 2010; Rahman, Tsukamoto, Rahman, Yoneyama, & Mostafa, 2013). The lignin-nitrogen ratio (L/N) drives the decomposition process; low ratios accelerate degradation and the release of nutrients to the soil (de Oliveira et al., 2016).

Until now, most studies on litter quantity and quality in tropical environments have been performed in mature and secondary forest or plantations, but the study of the role of scattered trees in pastures has been marginal. The objective of this research was to estimate the litterfall production during the dry season by scattered L. acapulcense and V. pennatula trees in pastures.

Materials and methods

The study was conducted in a pasture within a dry tropical forest ecosystem, in Paso Panal, Paso de Ovejas municipality, Veracruz, Mexico (19° 13' 14'' N and 96° 29' 5'' W at 167 m). The climate type is Aw0"(w) (i') g, classified as the driest of the warm and humid categories (Köppen classification, adapted by García, 2004). The average annual temperature is 25.2 °C, with an average annual precipitation of 909 mm. The landscape is highly fragmented due to agriculture and ranching, and only small patches of primary and secondary vegetation from TDF persists in the region.

Site description and tree selection

In a 10-ha pasture with silvopastoral management, 10 mature scattered L. acapulcense and 10 V. pennatula trees were selected. Even though previous studies stated low sample sizes (n = 5 litter traps) (Finotti, Freitas, Cerqueira, & Vieira, 2003) could be used in forest settings where canopy is more uniform, in this study traps were placed under 10 trees to compensate for any effect of heterogeneous canopies of scattered trees. Lysiloma acapulcense trees were 9.7 ± 1.5 m in height, and 45.9 ± 10.2 cm in diameter at breast height (± SE). Vachellia pennatula trees were 5.2 ± 0.5 m in height and 23.5 ± 4.5 cm in diameter at breast height (± SE).

Litterfall collection

To assess litterfall, 0.25 m2 circular traps were handmade with 1.0 mm nylon mesh fixed to iron frames, and protected with a 40 x 40 cm piece of cloth at the bottom to prevent the loss of small leaflets. Three traps were placed under each tree at 1 m aboveground, at midpoint between the base of the shaft and the drip zone (Figure 1). Trapped litter was collected every 30 days over two consecutive periods of natural litterfall in the area (December 2014 - May 2015 and November 2015 - April 2016). Lysiloma acapulcense and V. pennatula lose their leaves during the dry season, but L. acapulcense is partially deciduous (Camacho-Moreno et al., 2017). Litter was dried in a forced air oven at 65 °C for 48 h, and then separated into leaves, twigs, and fruits. Only leaf biomass was evaluated.

Figure 1 Location of litter traps under a Vachellia pennatula tree. 

Leaf litter lignin and nitrogen

Nitrogen and lignin content in the leaves of both tree species were measured in samples collected from four trees (of each species), selected at random during December 2014 to May 2015, and from four trees during November 2015 to April 2016. The total nitrogen content was quantified using the macro-Kjeldahl method (Association of Official Analytical Chemists [AOAC], 1980), and lignin content was determined using the fiber fraction method (Van Soest, Robertson, & Lewis, 1991).

Data analysis

To calculate litter quantity per unit area, the litter dry weight from each trap was divided by the area of the litter trap (0.25 m2), then multiplied by the number of traps (30) and extrapolated to one hectare and expressed as Mg of litter·ha-1·year-1 (1 Megagram [1 Mg] = 1 000 kg). Means from all variables were compared between species using a t-test in the GLIMMIX procedure in SAS (SAS Institute, 2010). Leaf-fall was modelled and three parameters were calculated for each species (the beginning, maximum, and end of leaf-fall) using an adjusted non-linear mixed model adapted by Beck, Atzberger, Høgda, Johansen, and Skidmore (2006) and Elmore, Guinn, Minsley, and Richards (2012) using the green-brown R package (The R Project for Statistical Computing, 2015). The model assumed that leaf-fall occurs following an annual cycle, but that these deciduous species lose their leaves mostly during the dry season (November to May).

The adjusted model was a double logistic function:

Lf=11+expc+a1-ta2-11+expc+a3-ta4

where, Lf is the proportion of leaf litter collected at time t, c is the random effect of tree, a 1 and a 3 are the midpoints of the ascending and descending sigmoid (respectively), and a 2 and a 4 are the parameters and represent the steepness of the curve. The sigmoid model more accurately describes the variation in leaf-fall throughout the year, reducing error estimation.

Results and Discussion

Scattered trees remaining after the fragmentation of TDF play important roles in maintaining ecosystem functions and services in modified environments (Manning, Fischer, & Lindenmayer, 2006) such as pastures. In this study, leaves comprised the largest fraction of the litter for both tree species (53 to 64 %), but fruits were also a large litter component (Figure 2). Although L. acapulcense yielded approximately 4.564 Mg·ha-1·year-1 of leaf litter, and V. pennatula 4.202 Mg·ha-1·year-1, the leaf litter quantities were similar (t = 0.9, df = 39, P = 0.37). These quantities from dispersed trees are significant because they represent the primary nutrient inputs to pastures. However, other authors have found greater litter quantities for other TDF species. For example, Gliricidia sepium (Jacq.) Kunth produced 10.42 Mg·ha-1, and Mimosa caesalpiniifolia Benth. 10.79 Mg·ha-1 in silvopastoral associations (de Oliveira et al., 2016). Pithecellobium dulce (Roxb.) Benth. released 13.83 Mg·ha-1·year-1, and Leucaena leucocephala (Lam.) de Wit produced 10.83 Mg·ha-1·year-1 in 10-year-old plantations (Ceccon, Sánchez, & Powers, 2015). In late successional forests of the dry tropics, Campo and Vázquez-Yañez (2004) also reported higher litter biomass (9.20 Mg·ha-1·year-1). Although only leaf litter was accounted for, lower amounts of litter assessed in this research may be explained by the fact that litter was collected only during the season of greater leaf-fall, discounting miscellaneous litter released year-round (de Queiroz et al., 2019). In addition, differences could arise from scattered trees being a part of simplified and low-biodiverse agricultural land units, as opposed to higher species-rich stands where different species have temporal and complementary dynamics of litterfall, leading to constant and higher litter supply in time and space (Huang, Ma, Niklaus, & Schmid, 2018). The litter biomass from both evaluated species are more similar to litter from early succession tropical forest stands (4.749 Mg·ha-1·year-1) (Aryal, De Jong, Ochoa-Gaona, Mendoza-Vega, & Esparza-Olguin, 2015), but there are few references of similar research to compare to these findings.

Figure 2 Composition of vegetative structures collected from the legume tree species studied in the pasture, during the dry season. 

Regarding leaf-fall timing, the parameters for the beginning and end of leaf-fall as well as maximum peak were significant (P < 0.0001) in the models for both tree species (Figure 3). The beginning of leaf-fall (start of season) was during autumn (on day 110 for V. pennatula, and day 122 for L. acapulcense). The time of maximum leaf-fall (peak of position) for V. pennatula was in December (day 157), and at the end of January (day 180) for L. acapulcense. The end of leaf-fall (end of season) took place towards the end of winter (February-March, day 205) for V. pennatula, and during spring (March-April, day 237) for L. acapulcense.

Figure 3 Annual model output of litterfall for the two legume tree species studied in the pasture, where the beginning, maximum and final point of leaf-fall are observed. 

The quantities of leaf litter produced by both tree species were similar, but the seasonal pattern of leaf-fall varied between the species. Although the start of leaf-fall appeared similar between the species, the maximum leaf-fall for V. pennatula occurred during December, while that for L. acapulcense occurred in January. The end of leaf-fall for V. pennatula was in February, while the leaf-fall for L. acapulcense ended between March and April. Yet, L. acapulcense did not completely lose its leaves because it is a semi-deciduous species (Camacho-Moreno et al., 2017). In environments with long seasonality, deciduous species completely lose their foliage, showing patterns of high leaf deposition towards the middle or end of the dry season (Kozovits et al., 2007). A similar pattern was observed for both species in this study, although the leaf-fall period was longer in L. acapulcense, as observed by Camacho-Moreno et al. (2017).

The concentration of nitrogen (1.49 and 1.94 %) and lignin (47.07 and 40.40 %) in leaf litter did not differ between L. acapulcense and V. pennatula (df = 7, P > 0.05) respectively, although the lignin-nitrogen ratio (L/N) was significantly higher in leaves from L. acapulcense (df = 7, P < 0.0001) than V. pennatula (Table 1). Oyun (2006) reported that some legume species such as L. leucocephala and G. sepium have leaves with high concentrations of nitrogen (4.5 and 4.8 %), intermediate concentrations of lignin (25.2 and 23.9 %), and low L/N ratios (5.6 and 5.0), respectively. As a result, their litter is considered of high quality. As suggested by the leaf-fall model, L. acapulcense retains its leaves for a long time, and may account for its higher L/N ratio. This would allow trees to mobilize nitrogen from the leaves, causing a greater imbalance in the ratio; however, the lower L/N ratio in V. pennatula litter is indicative of better litter quality and suggests that the leaf litter may be more labile than that from L. acapulcense. Leaf litter with a higher concentration of lignin degrades more slowly, partially reducing the nutrient release rate from the litter and the risk of loss by leaching (Krishna & Mohan, 2017). Thus, the combination of V. pennatula and L. acapulcense can be advantageous under agroforestry management. While V. pennatula leaves degrade faster, L. acapulcense leaves degrade slower, forming a temporary storage of carbon and nitrogen in the litter. However, more studies need to be conducted on other chemical components in the leaves of both species and on their degradation capacities.

Table 1 Chemical composition of leaf litter for the two legume tree species studied in pastures.  

Lysiloma acapulcense Vachellia pennatula t
Total N (%) 1.49 a 1.94 a 1.4
Lignin (%) 47.07 a 40.40 a 1.6
L/N ratio 31.5 a 20.8 b 10.2

Different letters in each row indicate statistically significant differences according to the t-test (P < 0.0001).

Scattered L. acapulcense trees are associated with pastures having long dry seasons, but the number of trees of this species declines due to fire and other management practices. On the other hand, V. pennatula trees are abundant in pasture lands regardless of fire and spraying for weed control performed by ranchers. Further efforts are needed to increase the cultural value of these TDF species for ranchers. Therefore, our research raises awareness of the importance of preserving the two species and enhancing the integration of these native trees into agroforestry management schemes. Still, further studies are required to explore the relationship between scattered tree density and litter production.

Conclusions

Litterfall from scattered trees is the primary source of nutrients for pasture soils. However, leaf litter deposited by scattered trees in pastures has been scarcely studied. This issue is particularly important because nutrient extraction from soils under grazing conditions is constant, with little or no nutrient inputs or amendments. Lysiloma acapulcense and Vachellia pennatula add important leaf litter quantities to pastures even as scattered trees. The different L/N ratios of these species suggest a complementary role in an agroforestry system because litter from V. pennatula has potential for more rapid decomposition and nutrient release while L. acapulcense litter might be a temporary carbon sink. Yet, there are gaps in the knowledge of ecological and biological traits of these species. Filling in these gaps will provide essential knowledge in the management of scattered trees in human modified landscapes; moreover, understanding the role of scattered trees in dry tropical pastures will help to improve agroforestry management.

Acknowledgments

We thank the site owner, Jorge Melchor Rivera, for his collaboration during the investigation, and to Colegio de Postgraduados for funding this study through the Línea Prioritaria de Investigación en Agroecosistemas Sustentables. Our gratitude to Maira and Ignacio Domínguez Lagunes, Eleonora Camacho Moreno, Diana Ríos Quiroz and Persia de Gante Ramírez for their valuable help with field work; to Dr. Jesús Jarillo Rodríguez and the Laboratorio de Nutrición y Forrajes in the Centro de Enseñanza, Investigación y Extensión en Ganadería Tropical of the Facultad de Medicina Veterinaria y Zootecnia, at the Universidad Nacional Autónoma de México for the chemical analyses. To Uduak Imeh for English revision and Yareni Perroni for their suggestions and comments.

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Received: December 30, 2019; Accepted: July 22, 2020

*Corresponding author: silvialopez@colpos.mx; tel.: +52 595 952 0200 ext. 3032.

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