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"Studies on i. | ' Neotropical Fauna — and Environment STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT Editors Joachim Adis, Max-Planck-Instiu fir Linnologie, AG Tropendkologie, Plin, Germany: Wolt Engels, Zoolagixches Unsstnr dey Universinit Tubingen, Tubingen, Germany: Brnstelosef Fittkau, fcking, Germany Managing Editor Anne Zillikens, Zoologisches Institut der Universinat Tabingen, Tabingen, Germany Hugo Campos, Instituto de Universidad Austral de Chile, Valdivia, Chile; Juliane Diller, Zool atssammlung, Munchen, Ger logy, Smithsonian Institute, Washington D.C., USA: Charles Heckman, Hamburg, Germany; Walter Hodi, Institut tur Zoologie der Universitit Wien, Austria; Vera Lucia Inperarrie-Fonseca Instituto de Biociéneias, Depto. Ecologia Geral, University of So Paulo, Si0 Paulo, Brazil; Hans-Wilkelm Koepke, Hamburg. Germany; Willian E. Magnusson, Departamento de Ecologia, INPA, Manaus. Brazil: Wolfgang Maier. Zoologisches Institut, Universitat Tubingen, Tubingen, Germany: Paul Miler. Institut fur Biogeographic, Universitit es Saarlandes, Ssarbricken, Germany: W-L. Overal, Muséu Emiftio Goeldi, Belém, Para, Brazil. Nelson Papavero, Instiuto de Estudos Avangados, Universidade de Sio Paulo, Sio Paulo, Brazil; David £. Pearson, Department of Zoology. Arizona State University, Tompe AZ, USA: Pedro Rexes-Castille, Instituto de Ecologia A.C., Veracruz. Mexico: Klaus Riede, Zoologisches lastitut der Universitit, Freiburs, Germany: Uivich Saint-Paul, Zenteum fir Marine ‘Tropendkologie, Bremen, Germany: Karl-Ludwig Schuchmann, Museum A. Konig, Bonn, Germany: José Willibatdo Thomé. Instituto de Biociéneias, PUC-RS. Porto Alegre, Brazil: Stefan Vidal. Institut fur Pflanzenkrankheit und Pfla venschutz, Hannover, Germany: Dieter Wirtmann, latitt fr Landwirtschattliche Zoologie und Bienenkunde. Univer sitit Bonn, Bona, Germany Publication programme 1996: Volume 31 (4 issues) Subscription price and ordering information Institutions: Df 367.00 including postage and handling. 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TERRESTRIAL FRurt CONSUMERS, ESPECIALLY THE WHITE-TAILED DEER (ODOCOILEUS 'S (SPONDIAS PURPUREA): IMPACT ON VIRGINIANUS) Salvador Mandujano! and Luis Enrique Martinez-Romero” Unstitute of Ecology, Xalapa, Veracruz, Mexico Biological Station Chamela, U.N.A.M., San Patricio, Jalisco, Mexico Received: 19 March 1996 Accepted: 9 April 1997 ABSTRACT In the deciduous forests ofthe tropical area of the Mexican Pacific coast, the tree Spondias purpurea is fruiting at the end of the dry season. The fruits are consumed by terrestrial vertebrates including the white-tailed deer (docoileus virginianus), If no free water sources are available, the deers’ liquid requirements may be met by “fruit consumption. By an exclusion experiment we were able to demonstrate that foraging chachatacas (Ortlis, poliocephala) increase the rare of frit fall about 5 times. KEYWORDS: Ontalis poliocephala, Odocoileus virginianus, Spondias purpurea, frait consumer, water supply INTRODUCTION In the tropical deciduous forests of the Pacific coast of Mexico, less than 20% of the annual rainfall oc~ curs during the six to seven month dry season (Bul: lock 1986). Rivers may have water for only a short period during the wet season (Cervantes & Mass 1988), These conditions cause considerable season- al changes in plant biomass (Bullock & Solis-Ma- anes 1990; Lott ef al 1987; Martinez-Yrizar et al 1992). At the end of the dry season, only a few trees are fruiting (Bullock & Solis-Magallanes 1990), in- cluding the red mombin tree, Spondias purpurea (Anacardiaceae) (Bullock & Solfs-Magallanes 1990). Correspondence to: S. Mandujano, Depto. Ecologia y Com: portamiento Animal, Instituto de Ecologia, Apdo. Postal 63, Xalapa 91000, Veracruz, México, Fax: +52-28-187808: e-mail: mandujan @ Mean fresh weight of fruits is 7.5 + 1.9 g and fruit production is estimated at 14.9 + 4.8 kg/ha (Mandu: jjano et al 1994), The water content of fruits at Chi mela was estimated, according to fresh and dry weight, to be 10 W/ha in 1991 (Mandujano & Gallina 1995b) ‘These fruits are eaten by mammals of 8 species: white- tailed deer (Odocoileus virginianus), collared pec cary (Pecari tajacu), coyote (Canis latrans), white-nosed coati (Nasua narica), gray fox (Urocy on cinereoargenteus), Virginia opossum (Didelphis virginiana), Pacific gray squirrel (Sciurus collie’) and deer mouse (Liomys pictus), and in addition by black iguanas (Ctenosaura pectinata), leaf-cutting ants (Atta sp.) and birds of 5 species: West Mexican chachalaca (Ortalis poliocephala), streak-backed oriole (leterus pustulatus}, yellow-winged cacique (Cacieus melanicterus), San Blas jay (Cyanocorax sanblassianus) and grayish saltator (Saltator coerw: lescens) (Berlanga, unpublished data). ‘Any factor influencing the rate of fruit fall from ted mombin trees would effect the consumption by terrestrial frugivorous animals (see Dubost 1984; Rob- inson & Eisenberg 1985; Smythe 1986), Here we report on an exclusion experiment undertaken to de- termine the fruit drop in dependance of arboreal con- sumers. MATERIALS AND METHODS ‘The study site Chamela is a 3300 ha nature reserve of the Biological Institute, Universidad Nacional Auténoma de México, and is located on the Pacific ‘coast in the state of Jalisco, Mexico (19° 30° N, 105° 03" W). Annual mean precipitation from 1977 to 1993 was 735 +210 mm. The rainy season extends from June or July to October. Annual mean temperature is 25°C (Bullock 1986), The dominant vegetation is tropical deciduous forest, with small patches of top- ical semideciduous forest in the larger valleys (Lott et al 1987). Within a forest area of about 50 ha, we selected 4 red mombin trees with over 50% ripe fruits. We ob- served each tree three times when arboreal consum- ces Were present and three times when they were absent during two-hour periods at 0800-1000, 1000 1200, 1200-1400 or 1600-1800 h. Only one tree was observed per period in random sequence. Ob- servations were made June 4-14, 1993, representing the second half of this fruiting season, Prior 10 observations the ground was cleared of fruits. The observer hid 20 m away from the respec~ live tree and counted the individuals of fruit forag- ing species. the number of fruits eaten, the number of fruits dropped, and the total foraging time per individual, In controls the observer was placed un- der the tree thus hindering the approach of any ani- mals The data were analyzed using the following sta- al tests: Kruskal-Wallis one-way analysis of tis variance on ranks, Spearman rank order correlations, for fruit fall in relation to the number and foraging time of chachalacas, and Mann-Whitney rank sum test (Zar 1984) for rate of fruit fall with and without chachalacas, S. MANDUIANO AND L.E. MARTINEZ-ROMERO RESULTS. In a total of 24 h of observation, 37 chachalacas and 1 Pacific gray-squirrel were recorded. The squirrel stayed in the tree for 20 min and dropped one fruit. Up to 8 chachalacas visited a particular red mombin tree within a 2 h period, with individual foraging time ranging from 2-20 min (X= 5.6 min/visit) and up to 3 visits per observation period. The number of Traits dropped was neither correlated with the number of chachalacas (r,= 0.20, P= 0.53) nor their foraging times (r,=-0.01, P= 0.97). The individual duration of foraging was not depending on the group size of chachalacas consuming fruits at the same time (r= 0.25, P= 0.43), When birds were present, the rate of fruits dropped was 5 fruits, compared to 1 fruit/h without birds (T= 206.0, P= 0.001). DISCUSSION ‘The relative importance of arboreal animals in mak- ing ted mombin fruits available on the ground can be better understood if we consider the weight, for- aging behavior, and population density of the differ- ent species. In the study area, chachalueas are the largest avian frugivores (body mass= 0.70.9 kg, Berlanga unpublished). They forage by jumping from branch to branch, often occur in family groups (Gur- rola unpublished) and have a relatively high popula- tion density (0.7 ind/ha, Ornelas er al 1993; Hutto 1994). Other fruiteating birds occurring sympatri- cally have no relevant influence on red mombin fruit fall, Groups of Pacific gray squirrels (body weight= 0.4-0.5 kg, Ceballos & Miranda 1986) were seen only in the semideciduous forest where red mombin trees are rare (Mandujano er al 1994), but single individuals were sometimes observed also in the dry forest (0.04 ind/ha, Mandujano 1996). Even by jump- ing from branch to branch the squirrels cause little fruit drop because of their low body weight. From 1989 to 1993, the population density of deer at Chamela varied between 0.11-0.14 deer ha (Man- diujano & Gallina 1995a) with a mean biomass of 3.3 kg/ha (Mandujano & Gallina 1995b). Under ther- moneutral conditions, daily water demand of indi- vidual white-tailed deer was estimated at $3119 ml/ kg of body weight (Hervert & Krausman 1986; Knox ‘etal 1969), Thus, water demand by the deer popula- FRUrT FALL Cause BY CHACHALACAS 3 tion at the end of the dry season was estimated at 0.17-30.39 litervha/day (Mandujano & Gallina 1995b). Because there is no source of free water during the dry season, sources like dew, leaves and fruits are important, of which red mombin fruits pro- ‘vide a spatially and temporally patchy resource (Man- dujano & Gallina 1995b), Based on a microhistolog- ical analysis of deer faeces and the kinds of regurgitated endocarps, in fact red mombin fruits constitute an important diet during the dry season (Mandujano et af 1994, Arceo, unpublished). The role of chachalacas in causing fruit fall from red mombin trees and the fruit consumption by the deers as influencing their population dynamics could be analyzed in more detail only by a long-term exclu- sion experiment. Our data show, however, that of the arboreal fruit consumers only chachalacas are likely to effect the availability of red mombin fruits on the ground. ‘Though we do not have quantitative data, we assume that chachalacas have similar impact on fruit fall of other trees since in February of 1992 we counted SO chachalacas feeding on fruits of a Ficus sp. tree. The birds dropped large quantities of fruits which were eaten by aherd of 9 collated peecaries, 7 white-tailed deer and 1 coati). Mandujano e7 al (1994) end Man- dujano & Gallina (1995b) suggested that Spondias purpurea fruits provide an important water source at the end of the dry season for several animal, prin pally the white-tailed deer. ACKNOWLEDGEMENTS: We thank J. Laborde, F. Gonzilez-Garcia, V. Rico-Gray, H. Gomez de Silva, F. Ornelas, G. Sinche7-Rojas, 8. Gall- ina, C. Dominguez and R. Tod Highsmith for critically reading of the ms. This work was supported by CONA- CYT project (PO20CCOR-903703). REFERENCES BULLOCK $ H (1986) Climate of Chamela, Jalisco, and trends in the South Coastal Region of Mexico. Arch Meteorol Geophys Bioctimatol 36: 29-16 BULLOCK SH, SOLIS-MAGALLANES J A (1990) Phe- nology of canopy trees of a deciduous forest in Mexi- co, Biotropica 22: 2-35 CEBALLOS G, MIRANDA A (1986) Los Mamiferos de Chamela, Jalisco, Instituto de Biologia, Universidad Nacional AutGnoma de México, México CERVANTES L, MASS M (1988) Relacién Iluvia-escur- rimiento en un sistema pequento de cuencas de selva ‘baja caducifolia, Ingenieria Hidradlica en México {enero-abril): 30-42 DUBOST G (1984) Comparison of the diets of frugivo- ous forest ruminants of Gabon. J Mammal 65: 298— 316 HERVERT J J, KRAUSMAN P R (1986) Desert mule deer use of water developments in Arizona. J Wildlife ‘Managem 50: 670-676 HUTTO R L (1994) The composition and social organiza- tion of mixed-species flocks in a tropical deciduous forest in western Mexico, Condor 96: 10-118 KNOX K L. NAGY J G, BROWN D R (1969) Water turnover in mule deer. J Wildlife Managem 33: 389— 393 LOTT E J, BULLOCK $ H, SOLIS: MAGALLANES J A, (1987) Florist diversity and structure of uptand and arroyo forests in coastal Jalisco. Biotropica 19: 228— 235 MANDUJANO 5 (1996) Densidad poblacional de 1a a dilla gris del Pacifico (Sciurus colliae) en un bosque tropical de Jalisco. Revista de Mastozoologia Mexica- nna 2: 89-95, MANDUJANO S, GALLINA § (1995a) Comparison of ‘deer censusing methods in tropical dry forest. Wildlife Soe Bull 23: 180-186 MANDUJANO S. GALLINA S (1995b) Disponibitidad de agua para e! Venado cola blanca en un bosque tropi- cal caducifolio en México. Vida Silvestre Neotropical 4: 107-118 MANDUJANO S, GALLINA 8, BULLOCK $ Hi (1992) Frugivory and dispersal of Spondias purpurea (Ana- cardiaceae) in a tropical deciduous forest in Mexico. Revta Biol Tropical 42: 105-112 MARTINEZ-YRIZAR A, SARUKHAN J, PEREZ-JIMEN- EZ. A, RINCON E, MAASS J M, SOLIS-MAGAL- LANES A, CERVANTES I. (1992) Above ground phy tomass of a tropical deciduous forest on the coast of Jalisco, Mexico. J Trop Ecol 8: 87-96 ORNBLAS F, ARIZMENDI M, MARQUEZ-VALDELA- MAR L, NAVARUO ML, BERLANGA A (1993) Var- ability profiles for fine transect bird censuses in a tropical Ary forest in Mexico. Condor 95: 422-441 ROBINSON J G, FISENBERG J F (1985) Group size and foraging habits ofthe collared peceary (Tayassu tajacw. J Mammal 66: 153-155 SMYTHE N (1986) Competition and resource partition ing in the guild of neotropical terrestrial frugivorous. mammals. Annu Rev Ecol System 17: 169-188 ZAR JH (1984) Biostatistical analysis, 2nd ed. Premtice Hall Ine, Englewood cliffs, NJ ims and Scope here is still a far from complete understanding of the complex ecosystems in the Neotropics, although they have been studied since the first expeditions of the old world naturalists Marcgrave, Humboldt, Spix, Darwin, Bates and Maller ‘The aims and scope of this journal are, besides taxonomic and zoogeographic surveys, analyses of animal commu hities and their relationship with biotic and abiotic environmental conditions. This includes both the fauna of terrestrial and fresh water ecosystems in the Neotropics. Contributions that represent original researeh and mini: reviews are welcome, Manuseripts Manuscripts, illustrations and all communications relative to them should be sent to the Managing Editor, Dr. Anne Zillikens, Zoologisches Institut, Universitit Tubingen, Auf der Morgenstelle 28, D-72076 Tiibingen, Ger many. 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