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Use of Space in Two Neighboring Groups of the Howler Monkey Alouatta Palliata Mexicana (Primates: Atelidae): Overlap and Home Range Size

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Use of Space in Two Neighboring Groups of the Howler Monkey Alouatta Palliata Mexicana (Primates: Atelidae): Overlap and Home Range Size THERYA, 2017, Vol. 8 (2): 91-97 DOI: 10.12933/therya-17-462 ISSN 2007-3364 Use of space in two neighboring groups of the howler monkey Alouatta palliata mexicana (Primates: Atelidae): overlap and home range size PAULO CÉSAR QUINTANA-MORALES1*, JORGE E. MORALES-MÁVIL1, MATEO ESCOBAR-ALIAGA1, AND ROCÍO BRAVO-XICOTENCATL1 1 Laboratorio de Biología de la Conducta, Instituto de Neuroetología, Universidad Veracruzana, Av. Luis Castelazo Ayala s/n, Colonia Industrial Ánimas, CP. 91190, Xalapa. Veracruz, México. Email: [email protected] (PCQM), [email protected] (JEMV), [email protected] (MEA), [email protected] (RBX). * Corresponding author The interaction between primate groups and the dynamics involved in the use of space are important factors affecting both competition for resources and intergroup dominance. In the present study we analyzed the differences in home range overlap and intergroup size, as well as the diversity of tree species within the home range, the Value Index for the key tree species, and the daily activity pattern of two neighboring groups of the howler monkey Alouatta palliata. From March 2002 to June 2003, the behavior and location of two groups of monkeys (G1, two adult males, four adult females and four infants; G2, six adult males, five adult females, one juvenile and three infants) were recorded in the same forest patch at Los Tuxtlas Biosphere Reserve, Veracruz, Mexico. Using the Minimum Convex Polygon (100 %), grid (100 m), and Kernel (adapted and fixed) methods, as well as transect sampling, significant differences (F1, 20 = 14.45, P = 0.001) were found in the home range size of the groups, being greater in G2. The habitat in the G2 home range had greater richness and diversity of tree species, as well as a higher Impor- tance Value Index for the key tree species (Ficus spp.) included in the howler monkey diet. On the other hand, significant differences between groups (Mann-Whitney U = 38, P < 0.05) were found only as regards locomotion behavior, likely due to a behavioral adaptation to optimize food resource use or search. The two groups displayed a similar use of food resources; however, there was a slight overlap in the home range between both groups with no direct aggression, likely because their habitat included sufficient space and food. Our data suggest that the reduced overlap may vary between neighboring groups because of factors such as group size. However, groups could weigh various factors to avoid home-range overlap. La interacción entre grupos vecinos de primates y la dinámica en el uso del espacio son factores importantes durante la competencia por los recursos y la dominancia intergrupal. En este trabajo analizamos las diferencias intergrupales con relación al tamaño y traslape del ámbito hogareño así como la diversidad de especies arbóreas, el Índice de Valor de Importancia de especies clave y el patrón diario de actividad de dos grupos vecinos de monos aulladores Alouatta palliata mexicana. De marzo 2002 a junio 2003 registramos la conducta y las ubicaciones de dos grupos de monos (formados, G1 por 2 machos adultos, 4 hembras adultas y 4 infantes, y G2 por 6 machos adultos, 5 hembras adultas, 1 joven y 3 infantes) en un mismo fragmento en la Reserva de la Biósfera Los Tuxtlas, Los Tuxtlas, Veracruz, México. Aplicando los métodos de Polígono Mínimo Convexo (100 %), Rejillas (100 m), y Kernel (adaptado y fijo) así como un muestreo por transectos se encontraron diferencias significativas (F1, 20 = 14.45, P = 0.001) en el ámbito hogareño de los grupos, siendo mayor el de G2. El hábitat en el ámbito hogareño del G2 tuvo una mayor riqueza y diversidad de especies arbóreas así como un valor más elevado en el Índice de Valor de Importancia de especies clave (Ficus spp.) en la dieta de los monos aulladores. Por otra parte, se encontraron diferencias significativas (Mann-Whitney U = 38,P < 0.05) sólo en la conducta de locomoción, posiblemente debido a un ajuste conductual para optimizar el aprovechamiento o la búsqueda de recursos alimentarios. Los recursos alimentarios utilizados por los dos grupos fueron similares; sin embargo, se registró un reducido traslape del ámbito hogareño entre los grupos y ningún tipo de agresión directa, lo cual puede deberse a que encuentran suficiente espacio y alimento en el há- bitat. Nuestros datos sugieren que el reducido traslape puede variar entre grupos vecinos a causa de factores como el tamaño del grupo. Sin embargo, es posible que los grupos decidan ponderar diversos factores a fin de evitar los traslapes de ámbito hogareño. Key words: competition; howler monkey; intergroup distance; overlapping home range; spatial interactions. © 2017 Asociación Mexicana de Mastozoología, www.mastozoologiamexicana.org Introduction Unlike a territory, where individuals must invest time The use of space is a key component of the ecological niche and energy to protect it (Kodric-Brown 1978; Amsler and for all animal species (Lopez-Darias et al. 2012). One of the Brown 2010), the HR can be shared either partially or essential parameters to quantify the use of space is the totally. In this sense, it is argued that the level of HR over- home range (Börger et al. 2008). The size and the internal lap in various primate species depends on both the condi- dynamics of the home range (HR) are influenced by the tions of the habitat and the physical characteristics of the environmental, biological, and social factors of each spe- species (Pearce et al. 2013). In the Neotropics, the howler cies. In the case of primates, key factors as group biomass monkey (Alouatta spp.) is considered a non-territorial spe- (Campos et al. 2014), group size (Shaffer 2014), habitat qual- cies (Milton 1980; Hopkins 2013) because the home range ity (Campera et al. 2014), vegetation type (Pebsworth et al. frequently overlaps. Hopkins (2013) has pointed out that 2012), seasonality (Pyritz et al. 2011), and population den- the separation between groups of howler monkeys seem sity (Glessner and Britt 2005). to be influenced by the hierarchical status of neighboring USE OF SPACE IN ALOUATTA PALLIATA groups. However, he makes no reference to the influence climate is warm and humid, with a dry season from March of habitat quality, access to resources and their availability. to May and a rainy season from June to February. Rainfall The objective of this work was to determine the size ranges from 3,500 to 4,000 mm per year (60 mm in the dri- of the home range and the extent of the spatial interac- est month), and the mean annual temperature is above tion between two groups of howler monkeys, based on 22 ºC (Soto 2004). Whithin this patch, Cristobal-Azkarate the overlap of their respective HR. We explored whether and Arroyo-Rodríguez (2007) estimated the presence of the tree species richness and diversity, and the size of the seven groups of Alouatta palliata mexicana; for this work, groups of monkeys, are associated with the use of space by two neighboring and potentially interacting groups were two neighboring groups of howler monkeys that inhabit selected, to obtain behavioral records. At the start of the a fragmented landscape in Los Tuxtlas Biosphere Reserve, study, the groups selected were the following: group 1 Veracruz, Mexico. (G1), 10 individuals (2 adult males, four adult females and four infants); group 2 (G2), 15 individuals (six adult males, Materials and Methods five adult females, a young and three infants). The num- ber of adults did not vary throughout the recording period; Study area and subjects. The study area is located in the however, new infants were born, so that at the end of the region of Los Tuxtlas, Veracruz, Mexico. This area is frag- recording period, G1 and G2 had four and three infants, mented and disturbed, with patches of primary vegetation respectively. The sex and age of individuals was deter- within a matrix of agricultural and livestock areas (Mendoza mined based on the classifications of Clarke (1990) and et al. 2005; Solórzano-García et al. 2012). The patch inhab- Domingo-Balcells and Veà-Varo (2009). ited by monkeys is an irregular-shaped polygon encom- passing 100 ha approximately, with connections that Location of groups and behavioral records. The location enable the displacement of groups to other forested areas of each group was recorded between March 2002 and June of similar size. It is located near the community of Montepio 2003 using a portable Global Positioning System (Garmin (18.63306° to 18.64778° N and -95.09556 to -95.07861°° W; model V plus). Groups were recorded through sightings in Figure 1), within Los Tuxtlas Biosphere Reserve, Veracruz, the morning (07:00 to 13:00 h) and evening (13:00 to 18:00 Mexico. The vegetation is evergreen forest with various h) which were alternated during 55 and 67 days (G1 and G2, degrees of disturbance. The canopy species include Lon- respectively). The difference in the recording days between chocarpus cruentus, Dussia mexicana, Nectandra ambigens, the two groups was due to the fact that G1 stabilized its Brosimum alicastrum, Ficus yoponensis, Ficus tecolutensis, HR in February 2003, while G2 did it in March; hence, the Pouteria sapota, Spondias radlkoferi and Bursera simaruba, latter required a greater number of days of follow-up and among others (Castillo-Campos and Laborde 2004). The a larger number of locations for the estimates. On aver- Figure 1. Location of the study area (box) where various groups of the howler monkey, Alouatta palliata, coexist in the region of Los Tuxtlas, Veracruz, Mexico, on the Gulf of Mexico coast. 92 THERYA Vol. 8 (2): 91-97 Quintana-Morales et al. Table 1. Estimates of the home range of two groups (G1 and G2) of the howling random factor and the group as a fixed factor.
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