International Journal of Biodiversity Science & Management Vol. 5, No. 3, September 2009, 115–131 Biodiversity and biogeographic significance of the Sierra Chinajá in Alta Verapaz, Guatemala: a first look Curan Bonhama*, Eduardo Sacayónb, Mercedes Barriosc, Sergio Perezd, Carlos Vásquez-Almazánd, Jose! Cajasd, Nicte! Ordoñezb, Enio Canoe and Fredy Archilaf Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 a College of Forestry and Conservation, University of Montana, Missoula, MT 59802, USA; bEscuela de Biologı́a, Universidad de San Carlos de Guatemala, Guatemala City, Guatemala; cCentro de Datos para la Conservación, Centro de Estudios Conservacionistas, USAC, Guatemala City, Guatemala; dMuseo de Historia Natural, Escuela de Biologı́a Universidad de San Carlos de Guatemala, Guatemala City, Guatemala; eLaboratorio de Entomologı́a, Universidad del Valle de Guatemala, Guatemala City, Guatemala; fEstación Experimental Familia Archila, Coban, Alta Verapaz, Guatemala A rapid biodiversity assessment was carried out in the Sierra Chinajá, Guatemala in order to support the conservation policies of the national agency for protected areas management. This study represents the first systematic account of the flora and fauna of the area. The floristic composition was surveyed using 21 Whitaker plots and non-systematic sampling of other vegetative strata. Bird communities were assessed using a system of point counts and mist nets. Bats were sampled with mist nets at 400 m and 600 m. For small terrestrial mammals, two transects with 90 traps each were established at low and high elevations. Reptiles and amphibians were collected through non-systematic walks through forest patches. Dung beetles were sampled with pitfall traps. A total of 309 plant species were found, including trees, orchids and bromeliads. The list of animal species includes 20 bats, four rodents, one marsupial, 110 birds, 24 reptiles, 14 amphibians, and 20 dung beetles. The results indicate that the Sierra Chinajá is an ecotone or transition zone from predominantly tropical lowland to a mix of montane environments. Keywords: biodiversity; rapid ecological assessment; ecotone; low elevation cloud forest Introduction In 1989, the Sierra Chinajá was declared by Guatemala’s protected areas management agency, the Consejo Nacional de Areas Protegidas (CONAP), as a special protection area. This is a temporary classification, which, by law, requires further technical studies in order to give the area operational management status. Although the Sierra Chinajá has been recognized as having natural resources of national importance that warrant protection, this area still has not been permanently classified under Guatemala’s system of protected area categories. Thus, it still lacks formal on-theground administration and management (Bonham et. al. 2008). For this reason, and the chaos resulting from a 30-year armed conflict, several indigenous communities have illegally settled in the Sierra Chinajá, which drastically affect its natural resources. The first step required by Guatemalan law to move the Sierra Chinajá from being an area of special protection (a ‘paper park’) to a functional protected area is the preparation of an ‘Estudio Te!cnico’ or technical study, as mandated by CONAP (1989). This study forms the baseline from which management plans and other multiple-use/concession plans can be formed. Part of the study involves an assessment of biodiversity. Prior to the research reported in this paper, no efforts had been made to document the biodiversity of the Sierra Chinajá. In order to assess the biological diversity of the area, a variety of rapid assessment methodologies were adapted to gather basic information. Rapid assessments of biodiversity, although incomplete due *Corresponding author. Email: curanbonham@hotmail.com ISSN 1745-1590 print/ISSN 1745-1604 online # 2009 Taylor & Francis DOI: 10.1080/17451590903223236 http://www.informaworld.com to limited time and financial constraints, are nonetheless useful in identifying relevant biodiversity values and potential management options (Sayre et al. 2000). The limitations of any study that is time-bound are obvious; however, by targeting certain taxa, through the use of indicator species, these studies can be used to tailor management plans to conserve areas with unique biogeographical value (Gaston and Blackburn 1995; Kerr et al. 2000; Oliver and Beattie 1993). In particular, dung beetles (family Scarabinae) are considered good ecological indicators because they are commonly found in distinct assemblages with specific biogeographical distributions (Favila and Halffter 1997). Additionally, amphibians are sensitive to habitat alteration and degradation due to climate change (Young et al. 2001). The need for high humidity and microhabitat niches (e.g. tank bromeliads, coarse woody debris and deep leaf litter) characteristic of mature forest limits the distributional range of many amphibian species, particularly tree and leaf frogs, to forest habitat. Therefore, these species may serve as ecological indicators (Pearman 1997). Furthermore, some amphibians are confined to a home range near their place of birth and therefore can be good indicators of local site conditions (Campbell and Vannini 1989). The information gathered through this study is currently being used by a local non-profit organisation (APROBASANK) to support CONAP in their efforts to change the status of the Sierra Chinajá to a more formal and exclusive 116 C. Bonham et al. category, and also to integrate the communities living in the area into its conservation and management. Study site Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 The Sierra Chinajá is located in the northern region of the Municipality of Chisec in the department of Alta Verapaz, Guatemala (Figure 1). It comprises 12,833 ha of karst mountains, ranging from 200–765 m. Replete with dissolution caves enshrouded by lowland and montane semi-deciduous tropical forest, this isolated mountain range marks the last massif between the Sierra Chamá and the expansive northern lowland limestone plateau of the Pe!ten. Its steep slopes give way to an interior upland plain that provides suitable lands for both agriculture and a variety of unique Figure 1. Location of Sierra Chinajá. highland species. Seventeen communities from the Qeqchı́ ethnic group (approximately 3220 inhabitants) live in the Sierra Chinajá. According to Holdridge (cited by Barrios 1995), the Sierra Chinajá belongs to the very wet sub-tropical warm forest life zone and is totally isolated from other mountain systems due to its geographic location and the historic pattern of land conversion. The climate of the area has been established from 9 years of observations at the nearest weather station in San Agustin Pe!ten, (16! 04’00’’N, 90! 26’20’’W) at 140 m, collected by the national weather forecast institution (INSIVUMEH). The mean annual temperature is 26! C, mean annual precipitation is 2252 mm and mean annual relative humidity is 83%. There are two seasons: the driest months (mean rainfall ,100 mm) occur International Journal of Biodiversity Science & Management from February to April; the wet season (mean rainfall .200 mm) occurs from June to December. Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 Materials and methods Systematic sampling was conducted between June and October 2005. Several taxonomic groups were selected based on their perceived potential as indicator species and biogeographic importance. Taxa were also chosen based on ease of data collection methods and availability of Guatemalan expert consultants and collaborators. This study focused on five groups: plants, avifauna, mastofauna, herpetofauna and entomofauna. Four study sites were selected for sampling based on their degree of accessibility, forest integrity and habitat type. These sites were located in undisturbed primary forest interspersed with a matrix of perennial and annual agricultural fields. The terrain possessed slopes between 25–35%, with well-drained, clayey soils. The study sites were: Site 1, Nueva Esperanza (400 m; Site 2, Mucbilha II (300 m); Site 3, Nueva Chinajá (615 m); and Site 4, Tzulul Qeqchi (750 m). However, not all taxa were sampled at all sites. The forest cover composition was inventoried in the areas surrounding each of the four study sites using five randomly located transects, with Whittaker plots of 20 x 50 m established every 500 m (Comiskey 1999). In total, 21 plots were established to assess the diversity and abundance of mature forest at upper elevations (.400 m). Species height, form class and diameter at breast height (DBH) of all trees .10 cm DBH were recorded. These parameters were analysed to estimate species importance on a per hectare basis. An importance value for each tree species was calculated based on the frequency, density and basal area (Matteucci and Colma 1982). The other plant strata were sampled non-systematically through a series of non-random transects and reconnaissance walks in various habitats at the study sites. All flowering and fruiting species were collected, catalogued, preserved, identified and deposited in national herbaria at the Center for Conservation Studies (CECON) and the Faculty of Biology at the University of San Carlos (BIGU). Point count methodology and mist nets were used to record bird species (USFS 2002). Thirty point counts were located along trails and roads in three sampling sites, Nueva Esperanza, Tzulul Qeqchi and Mucbilha. Each point was separated by 250 m, and 10 min was spent at each point to identify birds present using audio or visual means. All transects were begun at 05:30 h, the approximate time of sunrise, and ended no later than 10:00 h. In addition, an array of six mist nets, 7 · 2 m, was used to sample the cryptic understorey avian community. Species were identified according to Howell and Webb (1995). For bats, the trapping configuration consisted of five mist nets, each 12 m long, and one harp trap. The total effort at each of the three sites sampled (i.e. Tzulul Qeqchi, Mucbilha, Nueva Chinaja) was 100 h/net. These nets were opened shortly after sunset (18:30 h) and closed 4.5 h later (23:00 h), so as to be operable during peak feeding activity. 117 On capture, the time was noted, the individual was identified and reproductive condition determined using Medellı́n et al. (1997) and Reid (1997) before release. Small terrestrial mammals were surveyed at two sites, Nueva Chinajá and Mucbilha II. In each site, a transect was made, placing alternately a Sherman, Museum special and Victor Rat Trap every 10 m following trails through forest patches and cardamom plantations. Two additional Tomahawk traps were located in each transect. Ninety trapping stations were placed in Mucbilha II and 90 in Nueva Chinajá. Bait consisted of oats, peanut butter and raisins in a proportion of 2:1:1. Each trap was baited each morning at 06:00 h and was checked every evening. Animals collected were measured and identified with field guides and deposited at the mammal collection at the Natural History Museum of the University of San Carlos de Guatemala (MUSHNAT), where the identification was confirmed. The reptile communities were sampled through diurnal and nocturnal walks in forest patches and along foot trails in Mucbilha II and Nueva Chinajá. All specimens collected were identified and placed in the MUSHNAT collections. These walks lasted from 06:00 h to 10:00 h in the morning and from 18:00 h to 23:00 h in the evening, on five consecutive days in the wet season in September. The community of copronecrophagous beetle species was assessed in Nueva Esperanza, Mucbilha and Tzulul Qeqchi through a system of pitfall transects. In each study site, two transects were located along foot trails crossing forest patches. Each transect was 200-m long and had pitfall traps consisting of a 450-ml plastic container (11 cm tall · 11 cm diameter at the opening) placed every 20 m. The traps were dug into the ground, half filled with soil and horse dung in a 3:1 ratio, and partially covered with a lid that had a wedge removed, about 25% of the surface area, to allow entry, but also to complicate the exit of any individual lured into the trap. In order to avoid confusion, florescent flagging was used to mark the location of the traps. Traps were left for 24 h and recollected at 09:00 h on the following morning. Although this study was a rapid analysis and registered only a small portion of the total distribution of diversity and abundance of species in the Sierra Chinajá, it was designed so as to permit multivariate statistical analysis. A cluster analysis using the Simpson index was used to examine the evenness among biological communities and allow for an interpretation of the affinities of the three study sites. This analysis is presented in several dendrograms and rarefaction curves, which were generated using the program PAST ver. 1.81 (Hammer et al. 2001). Dendrograms were not generated for all groups of taxa as some of the sampling methods did not lend themselves well to statistical analysis or the data were too sparse. Results A total of 309 species of plants were found in the Sierra Chinajá: 77 trees, 141 shrubs, herbs and vines, 73 orchids and 18 bromeliads (Tables 1 and 2). The 10 species with 118 C. Bonham et al. Table 1. Plant species found in the Sierra Chinajá, according to Ve !liz, M. and Archila, F. Family Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 Acanthaceae Anacardiaceae Annonaceae Apocynaceae Araceae Araliaceae Arecaceae Asclepiadaceae Asteraceae Begoniaceae Bombacaceae Boraginaceae Bromeliaceae Species Aphelandra aurantiaca Aphelandra deppeana Aphelandra scabra Justicia sp. Mosquitoxylum jamaicense Astronium graveolens Metopium brownei Rhus striata Annona scleroderma Cymbopetalum penduliflorum Desmopsis stenopetala Guatteria anomala Aspidosperma cruentum Aspidosperma sp. Plumeria rubra Thevethia ahouai Tabernaemontana sp. Anthurium montanum Anthurium pentaphyllum var. bombacifollium Anthurium sp.1 Anthurium sp.2 Anthurium sp.3 Anthurium sp.4 Monstera sp. Spathyphyllum blandum Syngonium podophyllum Dendropanax arboreus Oreopanax obtusifollium Oreopanax sp. Chamaedorea elegans Chamaedorea sp. Chamaedorea tepejilote Chryosophila argentea Asclepias curasavica Ageratina sp. Melanthera nivea Neurolaena lobata Vernonia sp. Zexmenia salvinii Begonia manicata Begonia nelumbiifolia Begonia sp. 1 Begonia sp. 2 Begonia sp. 3 Begonia sp. 4 Begonia sp. 5 Begonia sp.6 Pseudobombax ellipticum Cordia gerascanthus Cordia alliodora Aechmea bracteata Androlepis skinneri Billbergia sp. Billbergia viridiflora Catopsis sp. Catopsis hahnii Catopsis (pending identification) Greigia sp. Pitcairnia sp. Pitcarnia wendlandlii Tillandsia sp. Tillandsia butzii (Continued ) Table 1. (Continued). Family 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 Burseraceae Caesalpinaceae Cactaceae Clusiaceae Combretaceae Comelinaceae Costaceae Cyclantaceae Cyperaceae Dennstaedtiaceae Dioscoreaceae Ebenaceae Euphorbiaceae Fabaceae Flacourtiaceae Gesneraceae Gutiferae Haemodoraceae Pteridophyta Species Tillandsia juncea Tillandsia matudae Tillandsia schiedeana Tillandsia valenzuelana Tillandsia bulbosa Vriesia heliconoides Bursera simaruba Protium copal Swartzia sp. Epyphillum oxypetalum Zygocactus sp. Calophyllum brasiliensis Clusia guatemalensis Clusia sp. Garcinia sp. Vismia camparaguay Terminalia amazonia Campelia zanonia Costus ruber Costus pulverulentus Asplundia microphylla Scleria sp. Dennstaedtia sp. Dioscorea sp. Dioscorea barttletii Diospyros sp. Acalypha costarricenses Acalypha glummifera Acalypha sp. Croton glabellus Dalechampia heteromorpha Disciphania calocarpa Euphorbia leucocephala Hieronyma alchorneoides Sebastiania longicuspis Acacia sp. Bauhinia divaricata Desmodium sp. Dialium guianense Lonchocarpus castilloi Lonchocarpus sp. Myroxylon balsamum Schizolobium sp. Swartzia sp. Swartzia standleyii Senna skinneri Senna sp. Vatairea lundellii Casearia sp. Allopectus vinaceus Vismia camparaguay Xiphidium caeruleum Adiantum radiata Adiantum sp.1 Adiantum sp.2 Anthryphylum onsiforme Asplenium sp. Blechnum schedianum Campyloneuron sp. Displazium plantaginifolium Elaphoglossum sp. Pleopeltis lanceolata Pleopeltis sp. Pleopeltis sp. (Continued ) International Journal of Biodiversity Science & Management Table 1. Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 Family 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 Table 1. (Continued). Heliconiaceae Labiaceae Lauraceae Liliaceae Malphigiaceae Maranthaceae Melastomataceae Meliaceae Menispermaceae Mimosaceae Moraceae Myristicaceae Myrsinaceae Myrtaceae Nyctaginaceae Ochnaceae Olecaceae Orchidaceae (Continued). Family Species Polypodium sp. Tectaria heraclifolia Thelypteris sp. Calathia unsigues Heliconia latispatha Heliconia sp. Salvia sp. Licaria capitata Licaria sp. Nectandra globosa Ocotea licaria Dracaena americana Mascagnia sp. Calathea allouia Clidemia petiolaris Clidemia sp. Conostegia xalapensis Miconia sp.1 Miconia sp.2 Topobea calicularis Topobea lavrigata Urera sp. Guarea glabra Swietenia macrophylla Trichilia glabra Trichilia sp.1 Trichilia sp.2 Hyperbaena Mexicana Cassia sp. Pithecelobium sp. Inga sp. Dorstenia lindleyana Ficus radula Brosimum alicastrum Coussopoa sp. Trophys racemosa Ficus sp. Virola guatemalensis Ardisia sp. Parathesis sp. Eugenia sp. Eugenia sp. Guapira sp. Ouratea lucens Olecaceae sp. Brassia caudate Campylocentrum scheidei Catasetum integerrimum Chysis bractescens Coelia bella Coelia sp. Corymborkis forcipigera Cranichis sp. Elleanthus capitatus Elleanthus caricoides Elleanthus graminifolius Elleanthus poiformis Schltr. Encyclia (Prosthechea) cochleata Encyclia (Prosthechea) pygmaea Encyclia asperula Encyclia sp. Epidendrum cf. veroscriptum Epidendrum isomerum Epidendrum nocturnum (Continued ) 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 254 119 Passifloraceae Piperaceae Species Epidendrum polyanthum Epidendrum rigidum Eurystyles sp. Gongora cassidea Goodyera sp. Hexadesmia imbricata Isochilus linearis Jacquiniella cobanensis Jacquiniella equitantifolia Lepidanthus pasanticus Maxillaria aciantha Maxillaria brunnea Maxillaria crassifolia Maxillaria densa Maxillaria meleagris Maxillaria muricata Maxillaria pulchra Maxillaria scorpioidea Maxillaria sp. Maxillaria uncata Maxillaria variabilis Mormolyca ringens Nidema boothii Notylia barkeria Oncidium oerstedii Ornithocephalus bicornis Pleurothalis grobyii Pleurothalis lewisii Pleurothalis pansamalae Pleurothalis sanchoi Pleurothalis segoviensis Pleurothallis yucatanensis Pleurothallis sp. Polystachia cerea Polystachya masayensis Ponera juncifolia Ponera striata Prostechea cochleata Prostechea fragans Prostechea pygmaea Sarcoglottis sp. Scaphyglothis lendyana Scaphyglothis sp. Scaphyglottis crurigera Sobralia fragrans Sobralia sp. Stanhopea aff. Oculata Stelis sp. 1 Stelis sp. 2 Stenorrhynchus coloratus Trichosalpinx violacea Trigonidium egertonianum Vittaria graminifolia Zootrophyon tribuloide Passiflora biflora Passiflora sp.1 Peperomia pellucida Peperomia sp.1 Peperomia sp.2 Peperomia sp.3 Piper aduncum Piper auritum Piper peltatum Piper sp. (Continued ) 120 C. Bonham et al. Table 1. (Continued). Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 Family 255 256 257 258 259 260 261 262 263 264 265 266 267 268 269 270 271 272 273 274 275 276 277 278 279 280 281 282 283 284 285 286 287 288 289 290 291 292 293 294 295 296 297 298 299 300 301 302 303 304 305 306 307 308 309 Poaceae Polygonaceae Polypodiaceae Rubiaceae Rutaceae Sapindaceae Sapotaceae Sauraureaceae Simaroubaceae Solanaceae Theaceae Tiliaceae Ulmaceae Urticaceae Verbenaceae Violaceae Vitaceae Vochisiaceae Zamiaceae Species Lasiacis divaricada Muhlenbergia sp. Olyra lalifolia Coccoloba sp.1 Polypodium sp. Alseis yucatanenses Cephaelis tomentosa Chiococca alba Guettarda combsii Hamelia patens Hamelia rovirosae Hoffmania sp. Morinda sp. Psychotria chiapensis Psychotria sp.1 Psychotria sp.2 Psychotria sp.3 Psychotria sp.4 Psychotria sp.5 Rondeletia buddleioides Rondeletia sp.1 Rondeletia sp.2 Simira salvadorensis Zanthoxylum belizense Paullinia sp. Cupania belizensis Cupania sp. Pouteria sp. Chrysophyllum mexicanum Manilkara zapota Sideroxylon capiri Saurauia sp. Picramnia sp. Simarouba glauca Cestrum nocturnum Lycianthes sp. Solanum sp. Witheringia sp. Ternstroema tepezapote Mortoniodendron sp. Luehea candida Heliocarpus donnellsmithii Trichospermum galliothi Trema micrantha Ampelocera hottlei Pilea sp. 1 Phenax hirtus Cornutia sp. Vitex gaumerii Rinorea guatemalensis Cissus sp. Vochysia guatemalensis Zamia monticola Ceratozamia robusta Zamia tuerckheimii highest importance values were: Terminalia amazonia, Bursera simaruba, Manilkara zapota, Pouteria amygdalina, Blomia pisca, Pouteria sp., Psidium sartorianum, Desmopsis stenopetala, Pseudobombax ellipticum and Lonchocarpus guatemalensis. A total of 77 tree species (578 individuals) were recorded in 21 plots. The standard error was 0.42 and the corresponding sampling error was 2.5%. A cluster analysis using the Simpson index was used to compare the similarity among the three sub-sites sampled: Nueva Esperanza, Tzulul Qeqchi and Mucbilha. These three sub-sites are compared in the Figures 2 and 3 using the diversity of trees, birds, beetles and the three groups combined as metrics for describing the degree of similarity among the sub-sites. The figures consistently group Nueva Esperanza and Mucbilha (sites A and C). Tzulul Qeqchi is the outlier, having an assemblage of species more specifically distributed to that particular area. The survey also revealed 110 bird species, 24 reptile species, 14 amphibian species, 20 bat species, five small terrestrial mammal species and 20 dung beetle species (Tables 3–8). Species accumulation curves were created to analyse and estimate the total diversity of all taxa sampled (Figure 3). None of the curves presented are approaching their asymptotes, which suggests that more sampling would provide a clearer understanding of total species diversity. The beetle and rodent curve is particularly steep, indicating significant diversity left to be recorded. Discussion Biogeographic status of Chinajá The Sierra Chinajá is structurally and floristically similar to both the limestone mountains to the east in Belize and to the west in Mexico (Breedlove 1981; Meerman and Matola, 2003). Because of the interface between lowland and montane habitats, many species that are commonly separated exist together in the Sierra Chinajá. The rapid nature of this investigation provides just a glimpse into the floristics and diversity of the Sierra Chinajá. The region may contain as many as 4,000 plant species (Martinez et al. 1994). The suitability of the habitat provided by the Sierra Chinajá to both lowland and highland species explains its species richness. Its biogeographic isolation is another factor that makes this mountainous forest likely habitat for endemic populations, which further contributes to its potential species richness. The majority of bird species (Table 3) are characteristic of the tropical lowlands of the Pe!ten, as evidenced by the presence of individuals of typical lowland families such as Furnariidae, Formicariidae, Cotingidae, Thraupinae (Stotz et al 1996; Howell and Webb 1995). Nevertheless, 16 species are characteristic of upland tropical forest avian communities. This is likely to reflect changes in habitat type due to the orographic uplift in this isolated mountain chain. In such ranges, ecological zones are often compressed, resulting in the distribution of high montane species at lower elevations (Grubb 1971; Whitmore 1998). This uplift has effectively isolated these 16 montane species as if they were on an island. Twenty bat species were recorded, of which Carollia sowelli was the most abundant (Table 4). The composition of the bat community of the Sierra Chinajá is similar to that of the bat communities of the Atlantic lowlands (Perez, S. International Journal of Biodiversity Science & Management Table 2. Tree strata data from Whitaker plots at Sierra Chinajá. Bonham, C. Common name Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 121 Canxan Palo jiote Chico zapote Silion Colorado Qeqitzol Zapotillo Paata che Cacaute Mapola Palo gusano Palo algodon Aqal Tamarindo Colay Tem Amate Ramon blanco Palo lagarto Chechen blanco Laurel de montana Jocote Chilecayote Manax Santa Maria Aguacatillo Balsamo Pok xik Izote Valerio blanco Copal pom Pomte Escobo negro Luin hembra Cansin Faisan Anonillo Anona de monte Valerio Colorado Medallo Scientific name Family V/ha Terminalia amazonia Bursera simaruba Manilkara zapota Pouteria amygdalina Blomia pisca Pouteria sp. Psidium sartorianum Desmopsis stenopetala Pseudobombax ellipticum Lonchocarpus guatemalensis Ochroma lagopus Eugenia spp. Dialium guianense Sickingia salvadorensis Sideroxylon capiri Ficus radula Brosimum alicastrum Zanthoxylum belizense Sebastiania longicuspis Cordia alliodora Spondias mombin Sapium spp. Hieronyma alchorneoides Calophyllum brasiliensis Nectandra globosa Myroxylon balsamum Coccoloba spp. Dracaena americana Aspidosperma megalocarpon Protium copal Combretaceae Burseraceae Sapotaceae Sapotaceae Sapindaceae Sapotaceae Myrtaceae Annonaceae Bombacaceae Fabaceae Bombacaceae Myrtaceae Fabaceae Rubiaceae Sapotaceae Moraceae Moraceae Rutaceae Euphorbiaceae Boraginaceae Anacardiaceae Euphorbiaceae Euphorbiaceae Clusiaceae Lauraceae Fabaceae Rubiaceae Liliaceae Apocynaceae Burseraceae Cordia gerascanthus Lonchocarpus castilloi Boraginaceae Fabaceae Guatteria anomala Annona scleroderma Aspidosperma cruenta Vatairea lundellii Annonaceae Annonaceae Apocynaceae Fabaceae 34.01 14.78 20.79 18.67 12.25 6.88 7.68 8.18 14.13 5.61 7.38 3.37 7.90 7.07 11.88 8.08 6.03 5.50 5.28 4.12 2.17 8.83 3.06 3.86 3.99 5.97 2.07 3.12 1.66 1.45 1.39 1.32 1.35 2.89 1.16 0.61 1.08 0.64 1.03 BA total F total 8.38 3.89 4.81 4.76 3.71 1.97 1.81 2.54 3.68 1.49 2.26 0.88 2.29 1.70 3.03 2.58 1.43 1.34 1.36 0.80 0.55 2.12 0.65 0.89 1.13 1.46 0.57 0.59 0.37 0.36 0.37 0.35 0.32 0.69 0.33 0.21 0.26 0.14 0.22 0.62 0.48 0.76 0.57 0.71 0.90 0.57 0.43 0.33 0.43 0.48 0.57 0.43 0.43 0.29 0.38 0.43 0.38 0.38 0.33 0.43 0.19 0.33 0.38 0.24 0.24 0.38 0.33 0.38 0.29 0.33 0.33 0.29 0.29 0.33 0.18 0.15 0.10 0.05 D/ha Bar Fr Dr IV N 8.57 10.24 2.94 1.89 15.07 18 36.19 4.75 2.27 7.98 15.01 76 18.10 5.88 3.60 3.99 13.47 38 21.90 5.82 2.70 4.83 13.35 46 20.48 4.54 3.36 4.52 12.42 43 20.00 2.41 4.27 4.41 11.09 42 22.38 2.21 2.70 4.94 9.85 47 17.14 3.11 2.04 3.78 8.92 36 10.95 4.49 1.56 2.42 8.47 23 17.14 1.82 2.04 3.78 7.64 36 11.43 2.76 2.27 2.52 7.56 24 15.24 1.08 2.70 3.36 7.14 32 9.52 2.80 2.04 2.10 6.93 20 10.95 2.08 2.04 2.42 6.53 23 4.76 3.70 1.37 1.05 6.13 10 5.24 3.16 1.80 1.16 6.11 11 8.57 1.75 2.04 1.89 5.68 18 8.10 1.64 1.80 1.79 5.23 17 7.14 1.67 1.80 1.58 5.04 15 8.57 0.97 1.56 1.89 4.43 18 7.14 0.68 2.04 1.58 4.29 15 2.86 2.59 0.90 0.63 4.12 6 7.62 0.80 1.56 1.68 4.04 16 5.24 1.08 1.80 1.16 4.04 11 5.24 1.38 1.14 1.16 3.67 11 3.33 1.79 1.14 0.74 3.66 7 3.81 0.69 1.80 0.84 3.34 8 4.76 0.72 1.56 1.05 3.33 10 4.76 0.45 1.80 1.05 3.30 10 6.67 0.44 1.37 1.47 3.29 14 5.24 0.46 1.56 1.16 3.18 11 5.24 0.42 1.56 1.16 3.14 11 5.71 0.40 1.37 1.26 3.03 12 2.86 0.85 1.37 0.63 2.85 6 3.33 0.41 1.56 0.74 2.71 7 2.86 0.25 0.85 0.63 1.74 6 2.38 0.31 0.71 0.53 1.55 5 1.90 0.17 0.47 0.42 1.07 4 0.95 0.27 0.24 0.21 0.71 2 Note: D/ha = Density per hectare; Bar = Relative basal area; F = Frequency; Fr = Relative frequency; BA = Basal area; IV = Importance value; V/ha = Volume per hectare. personal communication 2005). While most of these species have a lowland affinity, several species were characteristic of highland bat communities. Dermanura tolteca is a species that commonly inhabits mountains of medium elevation. D. tolteca shares the Sierra with two other species from the same genus, both characteristic of the lowlands: D. phaeotis and D. watsoni. This elevated diversity may suggest a relatively complex system of niche partitioning worthy of further study. Another indicator that Chinajá posesses elements of low montane cloud forest is the presence of Sturnira ludovici. This species, representative of medium-sized mountains, was found sympatrically with S. lilium, the sister species more typical of the lowlands (Perez et al. 2005). Of the five species of small terrestrial mammals found, the most important was Peromyscus mexicanus (Table 5). This is found in mountains of medium elevation and is absent in the adjacent lowland jungles of the Pete!n. This is a new distribution record according to the mammal collection of the MUSHNAT, reaching its most northern distribution in the Sierra Chinajá. The total of 14 species of amphibians and 24 of reptiles collected (Tables 6 and 7) is likely to be only a fraction of the total herpetofaunal diversity in the area. The majority of the Sierra Chinajá is generally below 600 m, thus the predominant herpetofaunal species are widely distributed in the Caribbean lowlands of Mesoamerica (Campbell and Vannini 1989). The presence of Agalychnis moreletii and A. callidryas is another example of lowland and montane habitat overlap. According to Campbell and Vannini (1989), A. moreletii has a relatively restricted distribution, occurring in mesic forests along streams flowing through mountainous regions and, therefore, it is less likely to be found than A. callidryas. It occurs from central Veracruz and northern Oaxaca, Mexico, southward on Atlantic slopes to Guatemala and Belize. In 122 C. Bonham et al. Trees C A Coleoptera B 0.9 0.93 0.8 0.9 0.7 Similarity Similarity 0.96 0.87 0.84 C 1 0.99 A B 0.6 0.5 0.81 0.4 0.78 0.3 0.75 0.2 0.72 0 0 0.4 0.8 1.2 1.6 2 2.4 2.8 3.2 3.6 4 Index value 0.4 0.8 1.2 1.6 2 2.4 2.8 3.2 3.6 Index value Birds B All 3 groups C A 0.95 0.95 0.9 0.9 0.85 0.85 Similarity Similarity Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 C 4 0.8 0.75 B 0.8 0.75 0.7 0.7 0.65 0.65 0.6 0.6 0.55 A 0.55 0 0.4 0.8 1.2 1.6 2 2.4 2.8 3.2 3.6 4 Index value 0 0.4 0.8 1.2 1.6 2 2.4 2.8 3.2 3.6 Index value 4 Figure 2. Dendrograms of the Simpson Index for taxa analyzed in the 3 sites sampled in the Sierra Chinajá, Guatemala from June to October 2005. (Sites: A-Nueva Esperanza (380 m), B-Tzulul Qeqchi (750 m), C-Mucbilha (400 m)). Guatemala, the species ranges from 500 m to 2130 m (Campbell and Vannini 1989). A. callidryas is commonly found in the Atlantic lowlands and foothills from southern Veracruz and northern Oaxaca, Mexico, southward through Mexico, Guatemala, Belize and eastern Panama. In Guatemala, this species ranges from near sea level to about 950 m (Campbell and Vannini 1989). The presence of Eleutherodactylus xucanebi is further evidence of the montane character of the Sierra Chinaja, as its distribution ranges between 500 and 1500 m. Thus, biogeographic isolation and biodiversity concentration could be occurring in the Sierra since it is an island of montane habitat surrounded by lowland forest habitat that is unlikely to support these highland species. Among the 20 species of dung beetle (Table 8), the presence of Copris laeviceps and Copris nubilosus suggests once again that Sierra Chinajá is an ecotone or area of transition, because these species are characteristic of two distinct biogeographic areas: the highlands of Alta Verapaz and the the lowlands of the Pe!ten. Copris nubilosus was first described in 2003 (Kohlmann et al. 2003) and until now was only reported from colder high-elevation cloud forests between 1350 and 1800 m, in the Sierra Cuchumatanes and Sierra de las Minas. Copris laeviceps is a characteristic lowland dung beetle recorded from sites in the Atlantic lowland forests of Izabal as well as the Pe!ten Plateau to the north (Kohlmann et al. 2003). Status of rare and endemic species One notable endemic population is that of Ceratozamia robusta (a member of the Cycad family). This Neotropical genus is distributed in mountainous parts of Mexico, Belize and Guatemala (Jones 1993). The species reaches its southernmost distribution in Guatemala and is a rare ornamental (Jones 1993). The conservation of rare endemics of market value is of principal importance to conservation and land management efforts. While six migratory bird species were recorded, including streaked flycatcher (Myodinastes maculates), northern waterthrush (Seiurus noveboracensis), black-and-white International Journal of Biodiversity Science & Management Beetles Birds 72 Taxa (95% confidence) 27 Taxa (95% confidence) 123 24 21 18 15 12 9 6 3 64 56 48 40 32 24 16 8 0 0 1 1.2 1.4 1.6 1.8 2 2.2 2.4 2.6 2.8 Samples 6 12 18 24 30 36 42 48 54 Samples Rodents Amphibians 8 18 Taxa (95% confidence) Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 Taxa (95% confidence) 7.2 6.4 5.6 4.8 4 3.2 2.4 1.6 16 14 12 10 8 6 4 2 0.8 0 0 5 1.2 1.8 2.4 3 3.6 4.2 4.8 5.4 6 6.6 Samples 10 15 20 25 30 35 40 45 Samples Reptiles Bats 27 Taxa (95% confidence) Taxa (95% confidence) 36 32 28 24 20 16 12 8 4 21 18 15 12 9 6 3 0 0 6 Figure 3. 24 12 18 24 30 36 Samples 42 48 54 4 8 12 16 20 24 28 32 36 Samples Species accumulation curves of taxa sampled in Sierra Chinajá, Guatemala from June to October 2005. warbler (Mniotilta varia), Kentucky warbler (Oporornis formosus), Canadian warbler (Wilsonia canadensis) and Baltimore oriole (Icteus galbula), more are expected to use the area as a wintering ground. It is worth mentioning that local residents recognized the horn-billed guan (Oreophasis derbianus), a rare endemic typical of highland forests, from illustrations presented to them and claimed it is found in the highest parts of the mountains. Similar results were recorded by Jolon (2003) in the nearby Candelaria Caves National Park. The presence of several unique Chiroptera species such as Mimon cozumelae, Trachops cirrhosus and Tonatia saurophila, restricted to well-developed mature lowland forests, is also an indicator of the healthy state of forests in the Sierra Chinajá (Fenton et al. 1992). Due to their habitat specificity, these species have been proposed as ecological indicators (Fenton et al. 1992). It is important to mention the presence of Diphylla ecaudata in Nueva Chinajá, an uncommon species in human-associated areas because it feeds on the blood of non-domesticated animals (Uieda 1992). The fact that two new Coleoptera species of the genera Canthydium and Onthophagus were found, as well as at least one new species of Passalid beetle (Passalidae) (Schuster, personal communication), suggests that the Sierra Chinajá may already have given 124 Table 3. C. Bonham et al. Bird species found in the Sierra Chinaja. According to Bonham, C. with collaboration from Stewart, S. and Tenes, D. Scientific name Amazilia candida Amazilia tzacatl Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 Amazona autumnalis Aratinga astec Arremon aurantrirostris Arremonops chloronotus Attila spadiceus Method of Relative Common name detection Site Sensitivity* abundance* Whitebellied emerald Rufoustailed hummingbird Redlored parrot Cntab* Conservation priority* Endemic* Habitat* PC, MN 1,2,3 M C LT 4 F1,F7 PC, MN 1,2,3 L C LT 4 F1E,F15,F7 1,2 M C LT 3 F1E,F15,F,F8 1,2,3 2 L M C F LT LT 4 4 F1E,F15 F1 2,3 L C LT 4 – M F LT 4 F1,F7,F4 – M F UT 4 F4,F1 M C LT 4 F1,F2 – M C HT 4 F1,F4,F15 – L C LT 4 N13,N6 – M F LT 4 F8,F7,F1E,F3 – 1,2,3 L M C F LT LT 4 4 F1E,F7,F8,F3, F1E,F8,F15,F7 – M F LT 4 2 L C UT 4 1,3 M F LT 4 1 – L L C C LT LT 4 4 N14,F7,F8,F15 A8,A6,A11,F14 – L C LT 4 A9,A6,A8,F14 M F LT 4 F1,F2,F15,F8 M F LT 4 F1,F4 3 L C LT 4 N14,N11,N1 2 – 1,2,3 L L L F/P C C HT LT LT 4 4 4 F1E,F4E,F7,F8 N16,N6,N1,N13 N14,N11 1,2,3 M F LT 4 F1,F15 2,3 L C/P LT 4 F1,F15,F8 2 H F LT 4 F1,F4,F7 2 H F LT 4 F1 2 L C LT 4 F1E,F15,F8 – L C LT 4 F3,F8,F15 – M C UT 4 F4,F1 – L C LT 4 A1,A2 PC Aztec parakeet PC Orangebilled PC, MN sparrow Greenbacked PC sparrow Brightrumped O attila Aulacorhynchus Emerald O prasinus toucanet Automolus Buffthroated PC ochrolaemus foliagegleaner Basileuterus Goldencrowned O culicivorus warbler Bubulcus (Ardeola) Cattle egret O ibis Buteo (Asturina) Grey hawk O nitidus Buteo magnirostris Roadside hawk O Campephilus Palebilled PC guatemalensis woodpecker Campylopterus Wedgetailed MN curvipenis saberwing Campylorhynchus Bandbacked PC zonatus wren Caryothraustes Blackfaced PC poliogaster grosbeak Cathartes aura Turkey vulture PC Ceryle torquata Ringed O kingfisher Chloroceryle Green kingfisher O Americana Chlorophanes spiza Green PC honeycreeper Columba Shortbilled O nigrirostris pigeon PC Columbina Ruddy talpacoti grounddove Contopus cinereus Tropical peewee PC Coragyps atratus Black vulture O Crotophaga Groovebilled ani PC sulcirostris Crypturellus Slatybreasted PC boucardi tinmous Cyanerpes cyaneus Redlegged PC honeycreeper Dendrocincla Ruddy MN homochroa woodcreeper Dendrocolaptes Barred PC certhia woodcreeper Dives dives Melodious PC blackbird Dryocopus lineatus Lineated O woodpecker Dysithamnus Plain antvireo O mentalis Egretta thula Snowy egret O 1 2 – Y Y F1E,F7,F15 F1,F15 F11,F4E,F1E Y F1,F15 (Continued ) International Journal of Biodiversity Science & Management Table 3. (Continued). Scientific name Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 125 Method of Relative Common name detection Site Sensitivity* abundance* PC 3 Elaenia flavogaster Yellowbellied elaenia Elanus leucurus Whitetailed kite O – Eupherusa e.eximia Stripetailed MN 2 hummingbird Euphonia goldi Olivebacked PC 1,2 euphonia Euphonia Yellowthroated PC 1,3 hirudinacea euphonia Falco rufigularis Bat falcon PC 2,3 Formicarius analis Blackfaced PC,MN 3 antthrush Galbula ruficauda Rufoustailed PC 2 jacamar Geotrygon Ruddy quaildove O – Montana Glaucidium Pygmy owl O – brasilianum Glyphorprhyncos Wedgebilled PC 3 spirurus woodcreeper Habia fuscicauda Redthroated PC 1,2 anttanager Habia rubica Redcrowned O – anttanager Heliothrix barroti Purplecrownd PC 2 fairy Henicorhina Greybreasted O – leucoprhrys woodwren Henicorhina Whitebreasted PC 1,2,3 leucosticte woodwren Hylomanes Tody motmot O – momotula Icterus Blackcowled PC 2 dominicensis oriole prosthemelas Icteus galbula Baltimore oriole O – bullockii Lanio aurantius Blackthroated O – shriketanager Laniocera Speckled PC 1 rufescens mourner Laterallus rubber Ruddy crake PC 2 Lepidocolaptes Streakheaded PC 3 souleyetti woodcreeper Leptopogon Sepiacapped PC,MN 3 amaurocephalus flyacatcher Leptotila casinii Greychested PC 2 dove Leptotila verreauxi Whitetipped PC 2 dove Leucopternis White hawk PC 3 albicollis Lipaugus unirufus Rufous piha MN 2 Malacoptila Whitewhiskered O – panamensis puffbird Manacus candei Whitecollared PC,MN 1,2 manakin Melanerpes Goldenfronted PC 1,2,3 aurifrons woodpecker Melanerpes Blackcheeked PC 2,3 pucherani woodpecker Microcerculus Nightingale wren O – philomela Cntab* Conservation priority* Endemic* Habitat* L C LT 4 N14,N11,F15E L M U/P U LT UT 4 4 N13,N14,N6 F1,F15 M F LT 4 L C LT 4 L M F C LT LT 4 4 L C LT 4 F1E,F M F LT 4 F1,F4,F7 L C LT 4 N1,N2,N14 M F LT 4 F1,F4 M F LT 4 F1E,F2,F15 H F LT 4 F1 M U LT 4 F1,F15 M C UT 4 F4 M F HT 4 F1,F4 H U HT 4 F1,F4 L F LT 4 L C UT 4 H F LT 3 M U/P LT 3 F1 L L F L LT LT 4 4 A1 F7,F8,F1, M F LT 4 F1,F15 M F LT 4 F7,F8,F1 L C UT 4 F7,F8,F15 H F LT 4 F1,F4,F7 M M F F LT LT 4 4 F1 F1,F15 M F LT 3 L C LT 4 N1,N2,F8 M C LT 4 F1,F15 H F HT 3 Y F1 F1E,F8,F15 Y Y F1E,F7E,F8E F1,F2 F1E, F15 F8,F7,F15 Y Y Y F1 FIE,F15 F1 (Continued ) 126 Table 3. C. Bonham et al. (Continued). Scientific name Dotwinged PC 1,2,3 antwren Ochrebellied MN 2 flycatcher Black and O – White warbler Bluecrowned PC 1,2 motmot Slatecolored PC 2 solitaire Myiobius Sulphurrumped PC 2 sulphureipygius flycatcher Myiozetetes similes Social flycatcher O – Myodinastes Streaked PC 2 maculates flycatcher Nyctidromus Pauraque PC 1,2 albicollis Oncostoma Northern PC,MN 2,3 cinereigulare bentbill Onychorhynchus Royal flycatcher PC 1 mexicanus Oporornis formosus Kentucky O – warbler Ortalis vetula Plain chachalaca PC 1,2,3 Pachyramphus Cinnamon PC 3 cinnamomeus becard Passerina Blueblack PC 2 cyanoides grosbeak Phaeochroa Scalybreasted O – cuvierrii hummingbird Phaethornis Little hermit PC,MN 1,2,3 longemareus Phaethornis Longtailed PC,MN 2,3 superciliosus hermit Piaya cayana Squirrel cuckoo O – Pionopsitta brownhooded O – haematotis Parrot Pipra mentalis Redcapped PC 1,2 manakin Piranga leucoptera Whitewinged PC 3 tanager Pitangus Great kiskadee PC 1,2 sulphuratus Psarocolius Montezuma O – montezuma oropendola Psarocolius wagleri Chestnutheaded PC 3 oropendola Psilorhinus morio Brown jay PC 1 Pteroglossus Collared aracari PC 1,2,3 torquatus Quiscalus Greattailed O – mexicanus grackle Ramphastos Keelbilled tucan PC 1,2,3 sulfuratus Ramphocaenus Longbilled PC 3 melanurus gnatwren Ramphocelus Scarletrumped PC 1,2 paserinii tanager Ramphocelus Crimsoncollared PC 1 sanguinolentus tanager Rhytipterna Rufous mourner O – holerythra Microrhopias quixensis Mionectes oleaginous Mniotilta varia Momotus momota Myadestes unicolor Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 Method of Relative Common name detection Site Sensitivity* abundance* Cntab* Conservation priority* Endemic* Habitat* M C/P LT 4 F1 M F LT 4 F1,F2,F15 L – MIGRANT 4 F1,F4,F15 M C LT 4 F1,F4,F15 M F UT 3 M F LT 4 F1,F8,F15 L L C C LT LT 4 4 F1E,F7E F1E,F15 L C LT 4 F1E,F15 L F LT 4 F1E,F7 H U LT 4 F1 – – MIGRANT – L L C F LT LT 4 4 M F LT 4 F1,F15 L C LT 4 F15,F1E M F LT 4 F1,F15 H C LT 4 F1,F4,F7 L M C F LT LT 4 4 F1,F7,F15 F1,F4 M F LT 4 F1 M F UT 4 F4,F1,F11 L C LT 4 F15,F8 M C LT 4 M F LT 4 F1,F15 L M F C LT LT 4 4 F8,F15 F1,F15 L C LT 4 N14,N13 M C LT 4 F1,F15 L FP LT 4 F1E,F15 L C LT 4 Y F1E,F15,N14 L F LT 4 Y F1E,F15 M F LT 4 Y F4,F11 – Y Y F1E,F8 F1E,F15 F1,F15 F1,F15 (Continued ) International Journal of Biodiversity Science & Management Table 3. (Continued). Scientific name Saltator atriceps Saltator coerulescens Saltator maximus Sclerurus guatemalensis Seiurus noveboracensis Sittasomus griseicapillus Sporophila torqueola Streptoprocne zonaris Tangara larvata Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 127 Tapera naevia Thamnophilus doliatus Thraupis abas Method of Relative Common name detection Site Sensitivity* abundance* Black-headed saltator Greyish saltator Buffthroated saltator Scalythroated leaftosser Northern waterthrush Olivaceous woodcreeper Whitecollared seedeater Whitecollared swift Goldenhooded tanager Striped cuckoo Barred antshrike Yellowwinged tanager Thraupis episcopus Bluegray tanager Thryothorus Spotbreasted maculipectus wren Tiaris olivacea Yellowfaced grassquit Tityra semifasciata Masked tatyra Tolmomyias Yellowolive sulphurescens flycatcher Trogon collaris Collared trogon Trogon Massena Slatytailed trogon Trogon violaceus Viloaceous trogon Turdus assimils Whitethroated thrush Turdus grayi Claycolored robin Tyrannus Tropical melancholicus kingbird Tyrannus savanna Forktailed flycatcher Veniliornis Smokybrown fumigatus woodpecker Volatinia jacarina Blueblack grassquit Wilsonia Canada warbler canadensis Xiphorhynchus Ivorybilled flavigaster woodcreeper Cntab* Conservation priority* Endemic* Habitat* PC 1,3 M F LT 3 F1E,F15 PC 1 L C LT 4 N14,N12 PC 2 L C LT 4 F1E,F15 H U LT 3 O – Y F1,F4 PC 2 M – MIGRANT 4 F1,F15,F14 PC 1 M C LT 4 F1,F2,F4 PC,MN 2,3 L C LT 4 N14,N1,N11 PC 1,2 L F LT 4 F4,F1,F15 PC 3 L C LT 4 F1E, F15 PC PC 2 1,3 L L C C LT LT 4 4 N14,N6,N11 N4,N11 PC 1,2,3 L C LT 4 F1E,F15,F8 O PC – 1 L L C F LT LT 4 4 F1E,F15 F1E,F15,F7 PC 1 L C LT 4 N14,N1 PC PC 1,2 1 M M C F LT LT 4 4 F1,F4,F15 F1,F4,F7 PC O 1,2 – M M C F LT LT 4 4 F1,F4,F2,F7 F1,F15 PC 1,2,3 M F LT 4 F1,F15 PC 1,2 M F UT 4 F4,F1,F7 PC 2 L C LT 4 F1E,F7,F15 PC 2 L C LT 4 F15,F8,F3 O – L C LT 4 N6,N7,N13 PC 1,2 L C UT 4 F4,F1,F15 PC 1,2,3 L C LT 4 N4,N6,N1 O – M – MIGRANT 4 F4,F15,F1 PC 1,2 M C LT 4 F1,F4,F7 Note: *According to Stotz et al. (1996) Center of Abundance: LT – Lower tropical (,500 m), LS – Lower subtropical (,500 m), HT -Hill tropical (500–900 m), UT – Upper tropical (900–1600 m), US – Upper subtropical (500–1600 m), MM – Middle montane (1600–2600 m), UM – Upper montane (.2600 m) Conservation Priority: 1 – Urgent, 2 – High, 3 – Medium, 4 – Low Endemic: Y = yes. rise to speciation, and raises the question to what extent it has served as a refuge through epochs of geologic and climatic change. Cross-site comparison The Simpson indices consistently group the study sites Mucbilha II and Nueva Esperanza, while separating out 128 C. Bonham et al. Table 4. Bat species found and their relative abundances according to Ordoñez, N. Pe !rez, S. and Cajas, J. Table 6. Reptiles registered in the Sierra Chinajá. According to Vásquez, C. with colaboration from Acevedo, M. Site Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 Species 1 Carollia perspicilliata 2 Carollia sowelli 3 Carollia brevicauda 4 Dermanura azteca 5 Dermanura phaeotis 6 Dermanura tolteca 7 Dermanura watsoni 8 Desmodus rotundus 9 Diphylla ecaudata 10 Glossophaga soricina 11 Mimon cozumelae 12 Myotis arbescens 13 Platyrrhinus helleri 14 Pteronotus helleri 15 Pteronotus parnelli 16 Sturnira ludovici 17 Tonatia saurophila 18 Trachops cirrhosus 19 Uroderma bilobatum 20 Sturnira lilium Total individuals captured Richness Mucbilhá II Nueva Chinajá Site Tzulul Qeqchi Total 3 3 7 10 10 17 10 3 2 2 2 2 1 3 5 5 6 1 1 2 2 1 1 1 3 1 1 3 1 1 1 2 2 2 1 2 2 1 1 4 8 38 22 4 67 4 13 6 20 2 1 Table 5. Small terrestrial mammals found in the Sierra Chinajá according to Ordoñez, N. and Pe!rez, S. Site Species 1 Peromyscus mexicanus 2 Oryzomys sp. 3 Sigmodon hispidus 4 Heteromys desmarestianus 5 Didelphis marsupiales Total individuals captured Mucbilhá II Nueva Chinajá 2 1 1 1 1 2 6 Tzulul Qeqchi. This may reflect the unique species assemblages present at Tzulul Qeqchi, which is located at a significantly higher elvevation. Tzulul Qeqchi and Nueva Chinajá represent the highest and best-conserved forests of the Sierra; correspondingly, the bat species richness is considerably higher. In Mucbilha II, the presence of vampire bat species, including the common widespread species Sturnira lilium, underscores the degree of environmental Species Mucbilha II 1 Ameiva festiva Wiegmann, 1834 2 Ameiva undulata 4 Lichtenstein, 1856 3 Atropoides nummifer Rüppel, 1845 4 Basiliscus vitattus 1 Wiegmann, 1828 5 Boa constrictor Linnaeus, 1758 6 Bothriechis schlegelii Berthold, 1846 7 Coniophanes fissidens 1 Günther, 1858 8 Dryadophis melanolomus Cope, 1868 9 Drymobius margaritiferus Schlegel, 1837 10 Eumeces sumicrasti Cope, 1866 11 Eumeces schwartzei 12 Imantodes cenchoa 2 Linnaeus, 1758 13 Leptodeira septentrionalis Günther, 1895 14 Leptophis aheatulla 1 Linnaeus, 1758 15 Ninia sebae Dume !ril, Bribon, and Dume !ril,1854 16 Norops biporcatus Dume !ril, Bribon, and Dume !ril, 1854 17 Norops capito Peters, 1863 18 Norops uniformis Cope, 7 1885 19 Pliocercus elapoides 1 Cope, 1860 20 Rhadinaea decorata Günther, 1858 21 Sceloporus teapensis Günther, 1890 2 22 Sibon sanniola Cope, 1867 23 Sphenomorphus cherriei Cope, 1867 24 Xenodon rabdocephalus 2 Wied, 1824 Total individuals captured 21 Richness n = 9 sp. Nueva Chinajá Total 5 5 4 3 3 1 1 1 1 1 1 1 1 1 1 1 1 1 2 4 1 1 1 2 2 3 3 1 1 4 11 1 1 1 4 4 2 3 3 2 35 56 n = 17 sp. n = 24 sp. perturbation. These species are commonly associated with zones of forest regeneration or areas that have been managed for agriculture (Fenton et al. 1992). Thus it is not surprising to find these species as well as D. rotundus, often associated with ranching, occurring sympatrically. P. mexicanus found in the upper elevations, was absent in the lower elevation sampling site of Mucbilha II and is International Journal of Biodiversity Science & Management Table 7. Amphibians found in the Sierra Chinajá, according to Vásquez, C. with help from Acevedo, M. Table 8. Dung beetles found in the Sierra Chinajá. According to E. Cano systematic entomology lab at Universidad del Valle. Site Especie Mucbilha II Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 1 2 3 4 5 6 Agalychnis callidryas Agalychnis morelleti Bufo valliceps 17 Eleutherodactylus alfredi Eleutherodactylus chac Eleutherodactylus laticeps 7 Eleutherodactylus psephosypharus 8 Eleutherodactylus xucanebi 9 Hyla microcephala 10 Leptodactylus labiales 1 11 Leptodactylus 2 melanonotus 12 Rana berlandieri 1 13 Rana vaillanti 1 14 Smilisca baudini 3 Total abundante 25 Richness n = 6 sp. Nueva Chinajá 1 1 3 1 3 2 Site Total 1 1 20 3 2 4 4 6 6 2 2 1 2 3 26 n = 10 sp. 129 1 1 6 51 n = 14 sp. also absent in the vicinities of Laguna Lachua National Park, a low-elevation tropical wet forest in the same region (Perez, S. personal communication 2005). Thus, P. mexicanus may now be isolated in the upper elevations of the Sierra Chinajá. At Tzulul Qeqchi (750 m), a community of seven species of hummingbirds (Trochilidae) included: Amazilia candida, Amazilia tzacatl, Phaethornis superciliosus, Phaethornis longuemareus, Campylopteris curvipennis, Eupherusa eximia and Phaeocroa cuvierrii. Based on the total number of species and the number of endemic species, Tzulul Qeqchi was identified as the most important site, with the greatest number of montane species. This is most likely because this site occupies the highest parts of the Sierra and has maintained a large degree of forest integrity. Conclusions An ecotone of elevated biodiversity This rapid assessment of biodiversity underscores the importance of the Sierra Chinajá as an ecotone between the Verapaz highlands and the Pe!ten lowlands, serving as habitat for both lowland and highland species. Due to its isolated position on the northern edge of the southern orographic uplift of Guatemala, the Sierra Chinajá may be one of the lowest elevation cloud forests in Mesoamerica where biogeographic speciation may be occurring. This assertion is warranted not only by the data, but also by the so-called Massenerhebung effect described by Grubb (1971), which causes the formation of montane forest conditions at lower elevations on narrow and isolated mountain ranges (Flenley 1995). Species 1 Ateuchus sp. 2 Bdeliropsis bowditchi 3 Canthon montanus 4 Canthydium sp.* 5 Copris laeviceps 6 Copris nubilosus 7 Deltochilum bowditchi 8 Deltochilum pseudoparile 9 Dichotomius agenor 10 Dichotomius satanas 11 Eurysternus angustulus 12 Eurysternus caribaeus 13 Ontherus mexicanus 14 Onthophagus sp.* 15 Onthophagus sp.1 16 Onthophagus sp.2 17 Onthophagus sp.3 18 Phanaeus endymium 19 Uroxys boneti 20 Uroxys micro Total abundance Mucbilha II Nueva Esperanza 50 4 9 3 31 3 3 1 8 Tzulul Qeqchi Total 8 8 54 1 1 30 3 6 3 70 3 6 1 1 6 6 8 15 8 1 1 2 2 1 1 2 2 1 1 1 1 2 78 1 1 3 3 2 189 53 58 Note: *Undescribed species (E. Cano, systematic entomology lab at Universidad del Valle). Many characteristic highland species are distributed at the upper elevational sites (.600 m) of Tzulul Qeqchi and Nueva Chinajá. These areas of lowland and highland ecosystem overlap are of primary conservation importance because of their elevated species richness. The presence of 16 typically highland avian species suggests a highland affinity of the avian community at higher sites. Not only is this pattern evident in birds, but also in dung beetles, where Copris nubilosus was found, a species previously reported only in Purulha, Alta Verapaz, a cloud forest habitat at approximately 1,200 m. The presence of this species in Sierra Chinajá is not only a new record, but also suggests the ecological importance of this mountain chain. The same pattern is true for Mastofauna, such as Dermanura tolteca, Sturnira ludovici and Peromyscus mexicanus, which are typical inhabitants of upland forests (Perez et al. 2005). The records of the leaf frogs Agalychnis moreletii, A. callidryas and Eleutherodactylus xucanebi are additional Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 130 C. Bonham et al. evidence of the highland nature and greater species richness of Sierra Chinajá as compared to adjacent lowland or highland ecosystems. While the majority of the Sierra Chinajá is habitat for predominantly lowland species, all the previously mentioned species are found only above 500 m and thus restricted to habitat that corresponds to less than 25% of the Sierra Chinajá; hence, populations of these species exist in an isolated highland community surrounded by lowland ecosystems. This suggests that they have persisted since the genetic flow between the mountainous areas of the Sierra Chama to the south was cut off. It therefore further suggests that speciation has been occurring within these populations for some time and that the probability of the existence of new endemic species is high. Nevertheless, some highland species may be able to migrate through, or seasonally occupy, lowland habitat, while some lowland species may be able to utilize highland habitat. The degree of habitat permeability or the ability of a species to freely pass from one side of this ecotone to the other warrants further study, and could contribute discoveries to biogeographic theory. The Sierra Chinajá illustrates an important ecological principle about the permeability of ecological boundaries and its effect on species distribution. The ecotone from predominantly tropical lowlands to a mix of montane environments is an enigma, and remains poorly understood by ecologists (Whitmore 1998). In the Sierra Chinajá, species assemblages change as one penetrates the interior highlands of the mountain massif. The change in elevation, and its corresponding effect upon species composition from predominantly lowland to predominantly highland communities, creates sympatric regions with habitat suitable for both lowland and highland species. As recognized by others (Wake 1987; McCain 2004, 2005), instead of increased competition leading to the exclusion of certain species, these data suggest that increased biodiversity results. lowlands has caused widespread dispersed settlement and resource extraction of this marginal area. For this reason, forest coverage is a mosaic of stand types in different successional stages, with only a few blocks of the most remote and inaccessible forest remaining in mature, well-developed cover. The upper elevational range (.500 m) of the Sierra Chinajá, where much of the unique biodiversity occurs, occupies very little surface area (,25% of the total range). Consequently, conservation of the highest forested slopes of the Sierra Chinajá should be a top priority for government agencies and conservation organisations. These forests provide habitat for many species dependent upon mature forests, as illustrated above. They also contain many valuable timber and nontimber forest resources (Salafsky et al. 1993). The exploitation of forest species has a long history in the lowlands of the Pe!ten (Schwartz 1990). Timber species such as Swietenia macrophylla and Cedrela odorata have been selectively logged from these areas since colonial times (Snook 1999). For this reason, and given the current global market for high-value tropical timber, these species are threatened throughout much of their range (Snook 1999). The Sierra Chinajá also provides habitat to many wellknown non-timber forest product species such as chicle (Manilkara sapota), allspice (Pimienta dioica), xate (Chamaedorea spp.), ramon (Brosimium alicastrum), sarsaparilla (Smilax sp.) and a variety of medicinal plants. The abundance and value of non-timber forest products in this region is a reason to assess development and conservation plans that take advantage of the sustainable management of these high-value species. The basic understanding of the biodiversity profile of this unique area provides a basis for government authorities to give it the attention it currently lacks and to grant it the management status that will assure its longterm continuity. Conservation importance Acknowledgements The Sierra Chinajá is the ecotone between the Lacandon Jungles and Verapaz Highlands, two areas of high biodiversity and conservation priorities for multi-million dollar biodiversity conservation finance programmes developed by international conservation organisations. Nevertheless, the Sierra Chinajá has not been the target of international conservation investments, despite possibly possessing as much or more ecological value as these two areas. Although the Sierra Chinajá is located in a remote zone with low population and has an average slope of ,30%, with shallow, rocky infertile soils, largely unsuited for agriculture, it has recently come under significant threat by land invasion, expansion of plantations and small-scale agriculture, illegal hunting and logging, unlicensed collection of ornamental plants, oil exploration and limestone mining. The inequitable distribution of agricultural lands in the more suitable adjacent We would like to thank the following institutions and personnel that helped to make this study possible: Manuel Vinicio Lopez, Teodoro Maas from APROBA-SANK, Reginaldo Reyes and the National Fund for Nature Conservation FONACON, Rosa Maria Chang from PROPETEN, Mario Veliz at the BIGU Herbarium, Students and faculty members from the vertebrate zoology department and the School of Biology from the Universidad de San Carlos who assisted in the fieldwork, Manuel Acevedo for his collaboration in the collection of herpetofauna, Daniel Tenes, who helped in bird surveys, Julio Morales and Miguel Flores from CECON Herbarium and the anonymous reviewers for providing invaluable and insightful input, which vastly improved this manuscript. References Barrios R. 1995. 50 Áreas de Intere!s Especial para la Conservación en Guatemala. CDC-CECON. The Nature Conservancy (TNC). Guatemala. 32. Downloaded By: [Bonham, Curan] At: 14:15 21 September 2009 International Journal of Biodiversity Science & Management Bonham C, Sacayon E, Tzi E. 2008. Protecting imperiled ‘‘paper parks’’: potential lessons from the Sierra Chinajá, Guatemala. Biodivers Conserv. 17(7):1581–1593. Breedlove DE. 1981. Introduction to the Flora of Chiapas. In Breedlove DE, editors. Flora of Chiapas. Part 1. San Francisco: California Academy of Sciences. p. 35. Campbell JA, Vannini JP. 1989. Distribution of amphibians and reptiles in Guatemala and Belize. West Found Vertebr Zool. 4(1). Comiskey J, Dallmeier E, and Mistry S. 1999. Protocolo de muestreo de vegetacion para la Selva Maya. Pp. 18–27 in: Carr III, A. and A.C. de Stoll (eds.) Monitoreo Biologico en la Selva Maya. U.S. Man and the Biosphere Program & Wildlife Conservation Society. PDF document, U.S.-MAB Program, Washington, 51 pp. CONAP. 1989. Decreto 4–89. Ley de Áreas Protegidas, reformas a la ley Decretos 18–89, 110–96 y 117–97, y reglamento. Congreso de la República de Guatemala. Guatemala. Favila ME, Halffter G. 1997. The use of indicator groups for mesuring biodiversity as related to community structure and function. Acta Zool Mex. 72:1–25. Flenley JR. 1995. Cloud forest, the Massenerhebung effect, and ultraviolet insolation. In: Hamilton LS, Juvik JO, Scatena FN, editors. Tropical montane cloud forests. Ecolgical studies (Vol. 110), New York: Springer-Verlag. pp. 150–155. Fenton M, Acharya L, Audet D, Hickey MBC, Merriman M, Obrist M, Syme D, Adkins B. 1992. Phyllostomid Bats (Chiroptera: Phyllostomidae) as Indicators of Habitat Disruption in the Neotropics. Biotropica. 24:440–446. Gaston K, Blackburn B. 1995. Mapping biodiversity using surrogates for species richness: macros-scales and New World birds. Proc R Soc Lond. 262:335–341. Grubb PJ. 1971. Interpretation of the ‘Massenerhebung’ Effect on Tropical Mountains. Nature. 229:44–45. Hammer Ø, Harper D, Ryan D. 2001. PAST: Paleontological statistics software package for education and data analysis. Palaeontol Electronica. 4:9. Howell S, Webb S. 1995. A guide to the birds of Mexico and Northern Central America. New York: Oxford University Press. p. 851. Jolon M. 2003. Biodiversidad en el Area del Las Cuevas de Candelaria: Fauna. Informe Final de Consultoria. FIPA. Guatemala. 25. Jones D. 1993. Cycads of the World. Washington (DC): Smithsonian Institution Press. pp. 255. Kerr J, Sugar A, Packer L. 2000. Indicator taxa, rapid biodiversity assessment, and nestedness in an endangered ecosystem. Conserv Biol. 14:1726–1734. Kohlmann B. Cano E. Delgado L. 2003. New species and records of Copris (Coleoptera: Scarabaeidae; Scarabaeinae) from Central America. Zootaxa. 167:1–16. Matteucci S, Colma A. 1982. Metodologı́a para el estudio de la vegetación. OEA. Serie biologı́a. Monografı́a. 22:168. Martinez E, Ramos C, Chiang C. 1994. Lista Floristica del la Lacandona, Chiapas, Mexico. Boletin de la sociedad Botanica de Mexico. 54:99–175. 131 McCain C. 2004. The mid-domain effect applied to elevational gradients: species richness of small mammals in Costa Rica. J Biogeogr. 31:19–31. McCain C. 2005. Elevational gradients in diversity of small mammals. Ecology. 8:366–372. Medellı́n RA, Arita HT, Sánchez O. 1997. Identificación de los murcie!lagos de Me !xico,clave de campo. Me!xico, DF: Talleres Offset Rebosán, S.A. p. 83. Meerman, JC, Matola S. (eds.), 2003. The Columbia River Forest Reserve: Little Quartz Ridge Expedition, A Biological Assessment. Columbia University Printing Services. 93 pp. Oliver I, Beattie A. 1993. A possible method for the rapid assessment of biodiversity. Conserv Biol. 7:562–568. Pearman P. 1997. Correlates of amphibian diversity in an altered landscape of amazonian ecuador. Conserv Biol. 11:1211–1225. Perez S, Castillo J, Echeverrria J, Masaya L, Jolon M. 2005. Las colleciones de mamiferos y las areas silvestres protegidas de Guatemala. Museo de Historia Natural. Universidad de San Carlos de Guatemala. Reid FA. 1997. A field guide to the mammals of Central America and Southeast Mexico. New York: Oxford University Press. p. 334. Salafsky N, Dugelby B, Terborgh J. 1993. Can extractive reserves save the rain forest? an ecological and socioeconomic comparison of nontimber forest product extraction systems in Peten, Guatemala, and West Kalimantan, Indonesia. Conserv Biol. 7:39–52. Sayre R, Roca E, Sedaghatkish G, Young B, Keel S, Roca R, Shepard S. 2000. Nature in focus: rapid ecological assessment. the nature conservancy. Washington DC: Island Press. Schwartz NB. 1990. Forest society: a social history of Peten, Guatemala. Philadelphia: University of Pennsylvania Press. Snook, L. 1999. Aprovechamiento sostenido de caoba (Swietenia macrophylla King) de las selvas de la penı́nsula de Yucatán, Me!xico: pasado, presente y futuro. In Primack, RB; Bray, D; Galletti, HA; Ponciano, I. eds. La Selva Maya: conservación y desarrollo. Me!xico, DF., Siglo XXI. p. 89–119. Stotz D, Fitzpatrick J, Parker T, Molkovits M. 1996. Neotropical birds: ecology and conservation. Chicago: The University of Chicago Press. p. 480. Uieda W. 1992. Periodo de Atividade Alimentar e Tipos de Presa dos Morcegos Hematófagos No Sudeste do Brasil. Rev Brasil Biol. 52(4):563–573. United States Forest Service. 2002. Bird Monitoring Techniques. Pacific Southwest Reesearch Station. General Technical Report: PSW-GTR-144 Wake DB. 1987. Adaptive radiation of salamanders in Middle American cloud forests. Annals of the Missouri Botanical Garden. 74(2):242–264. Whitmore T. 1998. An introduction to tropical rain forests. Oxford: Oxford University Press. p. 282. Young B, Lips K, Reaser J, Ibanez R, Salas A, Cedeno J, Coloma L, Ron S, La Marca E, Meyer J, Munoz A, Bolanos F, Chaves G, Romo D. 2001. Population declines and priorities for amphibian conservation in Latin America. Conserv Biol. 15:1213–1223.